project report submitted to the compatible
TRANSCRIPT
ldquoEFFECT OF LOW MOISTURE STRESS ON MORPHO-PHYSIOLOGICAL CHARACTERS AND YIELD RESPONSES
OF TWO PULSE CULTIVARSrdquo
Project report submitted to the
Department of Environmental Science
School of Science and Technology
Tezpur University
For the partial fulfilment of the degree
Master of ScienceIn
Environmental Science
Submitted by-Velentina Das
MSc4th semesterRoll No ESE10010
Department of Environmental scienceTezpur University
TezpurAssam
1
Department of Environmental Science
Tezpur University
This is to certify that the project report entitled ldquoEffect of Low-Moisture Stress on Morpho-
physiological Parameters and Yield Responses of two Pulses Cultivarsrdquo submitted to the
department of Environmental Science Tezpur University for the partial fulfilment of the
requirements for the award of the degree of Master of Science in Environmental Science is a
record of original work done by Ms Velentina Das under the supervision of Dr(Mrs) Nirmali Gogoi
in the Department of Environmental Science Tezpur University Assam
Head of the Department Supervisor(Dr RR Hoque) (Dr Nirmali Gogoi)
2
Department of Environmental Science Tezpur University Assam
I Velentina Das hereby declare that the project report entitled ldquoEffect of low moisture stress on Morpho-physiological Parameters and Yield Responses of Two Pulse Cultivarsrdquo submitted to Tezpur University in partial fulfilment of the requirement for the award of the degree of Master of Science in Environmental Science is a record of original work done during the autumn semester 2011 in the department of Environmental Science Tezpur University Assam
3
Velentina Das
I at the very onset would like to express my special thanks of gratitude to my Supervisor DrNirmali
Gogoi marsquoam Assistant Professor Department of Environmental Science Tezpur University for
providing ample of support valuable suggestion guidance and encouragement and cooperation at all
stages of my Project work She being an ideal person was always been my inspiration in every step
I offer my sincere thanks and gratitude to Professor KKBaruah Sir Tezpur University for his awful
lectures delivered in the regular classes which made me know my Project Topic more familiar before
head And along with his constant assistance suggestions guidance and co-operation and
encouragement during the entire course of project work
I would like to take the opportunity to thank and express my gratefulness to Dr KP Sarma Sir for
his guidance and providing me all the necessary fulfilments for the completion of the project work
I also convey my cordial thanks to DrRRHoque sir Head of the Department Department of
Environmental Science Tezpur University for giving me the golden opportunity to do this wonderful
project which not only helped me in doing a lot of Research but I explored many such events which
has been confined only to book and my limit of understanding was all based on theoretical
perspectives But after this project I could rather say that what I have gained is no more theoretical
now All I know is now practical
I also express and deliver my heartfelt Thanks to all the other Faculty members of Department of
Environmental Science- DrAKDas sir DrAshalata Devi marsquoam DrSSBhattacharya sir
4
DrMKumar sir Sumi Handique marsquoam for kind help and co-operation and encouragement during
the entire course of Project work
I would like to give my heartiest Thanks to Ms Bhaswatee Baroowa (Research Scholar) for being so
kind and helpful all throughout my Project She has been with me just like my elder sister in all phase
and makes me homely and giving moral support in every step
I express my cordial Thanks to my other Research Scholars who been giving constant advice
valuable suggestions and co-operation
Lastly but not least I would also like to thank my Parents My Local Guardian Dipankar Kalita
Department of Food Processing Technology Tezpur University and all my class-mate My Hostel
mates and my Roommate for being so humble and kind and along with helped me a lot in finishing
this project within the limited time
I am making this project not only for marks and partial fulfilment of the syllabus but its and handy
full aid to increase my knowledge and enlarge the circumference of my thoughts in every field
ldquoTHANKS AGAIN TO ALL WHO HELPED MErdquo
Abstract
Scarcity of water is a severe environmental constraint to plant productivity Drought-induced
loss in crop yield probably exceeds losses from all other causes since both the severity and
duration of the stress are critical Crop growth and productivity is adversely affected by
naturersquos wrath in the form of various biotic and abiotic stresses which adversely affects crop
growth and yield Therefore the present study was conducted with two major pulse crops
green gram (Vigna radiata L) and black gram (Vigna mungo L) in the sight to gain
information on various physiological changes taking place in the plants under low moisture
stress and its subsequent relation to plant growth development and yield Positive correlation
was found in plant height and leaf number and leaf area in both the pulses Regarding yield
both the pulses showed positive correlation with respect to soil moisture content The study
revealed that C1(T9) black gram is resistant than green gram against short-term moisture
stress And further it could be find out that the C3 (Pratap) of green gram has severe impact
on the moisture stress
5
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Department of Environmental Science
Tezpur University
This is to certify that the project report entitled ldquoEffect of Low-Moisture Stress on Morpho-
physiological Parameters and Yield Responses of two Pulses Cultivarsrdquo submitted to the
department of Environmental Science Tezpur University for the partial fulfilment of the
requirements for the award of the degree of Master of Science in Environmental Science is a
record of original work done by Ms Velentina Das under the supervision of Dr(Mrs) Nirmali Gogoi
in the Department of Environmental Science Tezpur University Assam
Head of the Department Supervisor(Dr RR Hoque) (Dr Nirmali Gogoi)
2
Department of Environmental Science Tezpur University Assam
I Velentina Das hereby declare that the project report entitled ldquoEffect of low moisture stress on Morpho-physiological Parameters and Yield Responses of Two Pulse Cultivarsrdquo submitted to Tezpur University in partial fulfilment of the requirement for the award of the degree of Master of Science in Environmental Science is a record of original work done during the autumn semester 2011 in the department of Environmental Science Tezpur University Assam
3
Velentina Das
I at the very onset would like to express my special thanks of gratitude to my Supervisor DrNirmali
Gogoi marsquoam Assistant Professor Department of Environmental Science Tezpur University for
providing ample of support valuable suggestion guidance and encouragement and cooperation at all
stages of my Project work She being an ideal person was always been my inspiration in every step
I offer my sincere thanks and gratitude to Professor KKBaruah Sir Tezpur University for his awful
lectures delivered in the regular classes which made me know my Project Topic more familiar before
head And along with his constant assistance suggestions guidance and co-operation and
encouragement during the entire course of project work
I would like to take the opportunity to thank and express my gratefulness to Dr KP Sarma Sir for
his guidance and providing me all the necessary fulfilments for the completion of the project work
I also convey my cordial thanks to DrRRHoque sir Head of the Department Department of
Environmental Science Tezpur University for giving me the golden opportunity to do this wonderful
project which not only helped me in doing a lot of Research but I explored many such events which
has been confined only to book and my limit of understanding was all based on theoretical
perspectives But after this project I could rather say that what I have gained is no more theoretical
now All I know is now practical
I also express and deliver my heartfelt Thanks to all the other Faculty members of Department of
Environmental Science- DrAKDas sir DrAshalata Devi marsquoam DrSSBhattacharya sir
4
DrMKumar sir Sumi Handique marsquoam for kind help and co-operation and encouragement during
the entire course of Project work
I would like to give my heartiest Thanks to Ms Bhaswatee Baroowa (Research Scholar) for being so
kind and helpful all throughout my Project She has been with me just like my elder sister in all phase
and makes me homely and giving moral support in every step
I express my cordial Thanks to my other Research Scholars who been giving constant advice
valuable suggestions and co-operation
Lastly but not least I would also like to thank my Parents My Local Guardian Dipankar Kalita
Department of Food Processing Technology Tezpur University and all my class-mate My Hostel
mates and my Roommate for being so humble and kind and along with helped me a lot in finishing
this project within the limited time
I am making this project not only for marks and partial fulfilment of the syllabus but its and handy
full aid to increase my knowledge and enlarge the circumference of my thoughts in every field
ldquoTHANKS AGAIN TO ALL WHO HELPED MErdquo
Abstract
Scarcity of water is a severe environmental constraint to plant productivity Drought-induced
loss in crop yield probably exceeds losses from all other causes since both the severity and
duration of the stress are critical Crop growth and productivity is adversely affected by
naturersquos wrath in the form of various biotic and abiotic stresses which adversely affects crop
growth and yield Therefore the present study was conducted with two major pulse crops
green gram (Vigna radiata L) and black gram (Vigna mungo L) in the sight to gain
information on various physiological changes taking place in the plants under low moisture
stress and its subsequent relation to plant growth development and yield Positive correlation
was found in plant height and leaf number and leaf area in both the pulses Regarding yield
both the pulses showed positive correlation with respect to soil moisture content The study
revealed that C1(T9) black gram is resistant than green gram against short-term moisture
stress And further it could be find out that the C3 (Pratap) of green gram has severe impact
on the moisture stress
5
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Department of Environmental Science Tezpur University Assam
I Velentina Das hereby declare that the project report entitled ldquoEffect of low moisture stress on Morpho-physiological Parameters and Yield Responses of Two Pulse Cultivarsrdquo submitted to Tezpur University in partial fulfilment of the requirement for the award of the degree of Master of Science in Environmental Science is a record of original work done during the autumn semester 2011 in the department of Environmental Science Tezpur University Assam
3
Velentina Das
I at the very onset would like to express my special thanks of gratitude to my Supervisor DrNirmali
Gogoi marsquoam Assistant Professor Department of Environmental Science Tezpur University for
providing ample of support valuable suggestion guidance and encouragement and cooperation at all
stages of my Project work She being an ideal person was always been my inspiration in every step
I offer my sincere thanks and gratitude to Professor KKBaruah Sir Tezpur University for his awful
lectures delivered in the regular classes which made me know my Project Topic more familiar before
head And along with his constant assistance suggestions guidance and co-operation and
encouragement during the entire course of project work
I would like to take the opportunity to thank and express my gratefulness to Dr KP Sarma Sir for
his guidance and providing me all the necessary fulfilments for the completion of the project work
I also convey my cordial thanks to DrRRHoque sir Head of the Department Department of
Environmental Science Tezpur University for giving me the golden opportunity to do this wonderful
project which not only helped me in doing a lot of Research but I explored many such events which
has been confined only to book and my limit of understanding was all based on theoretical
perspectives But after this project I could rather say that what I have gained is no more theoretical
now All I know is now practical
I also express and deliver my heartfelt Thanks to all the other Faculty members of Department of
Environmental Science- DrAKDas sir DrAshalata Devi marsquoam DrSSBhattacharya sir
4
DrMKumar sir Sumi Handique marsquoam for kind help and co-operation and encouragement during
the entire course of Project work
I would like to give my heartiest Thanks to Ms Bhaswatee Baroowa (Research Scholar) for being so
kind and helpful all throughout my Project She has been with me just like my elder sister in all phase
and makes me homely and giving moral support in every step
I express my cordial Thanks to my other Research Scholars who been giving constant advice
valuable suggestions and co-operation
Lastly but not least I would also like to thank my Parents My Local Guardian Dipankar Kalita
Department of Food Processing Technology Tezpur University and all my class-mate My Hostel
mates and my Roommate for being so humble and kind and along with helped me a lot in finishing
this project within the limited time
I am making this project not only for marks and partial fulfilment of the syllabus but its and handy
full aid to increase my knowledge and enlarge the circumference of my thoughts in every field
ldquoTHANKS AGAIN TO ALL WHO HELPED MErdquo
Abstract
Scarcity of water is a severe environmental constraint to plant productivity Drought-induced
loss in crop yield probably exceeds losses from all other causes since both the severity and
duration of the stress are critical Crop growth and productivity is adversely affected by
naturersquos wrath in the form of various biotic and abiotic stresses which adversely affects crop
growth and yield Therefore the present study was conducted with two major pulse crops
green gram (Vigna radiata L) and black gram (Vigna mungo L) in the sight to gain
information on various physiological changes taking place in the plants under low moisture
stress and its subsequent relation to plant growth development and yield Positive correlation
was found in plant height and leaf number and leaf area in both the pulses Regarding yield
both the pulses showed positive correlation with respect to soil moisture content The study
revealed that C1(T9) black gram is resistant than green gram against short-term moisture
stress And further it could be find out that the C3 (Pratap) of green gram has severe impact
on the moisture stress
5
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
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48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
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Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Velentina Das
I at the very onset would like to express my special thanks of gratitude to my Supervisor DrNirmali
Gogoi marsquoam Assistant Professor Department of Environmental Science Tezpur University for
providing ample of support valuable suggestion guidance and encouragement and cooperation at all
stages of my Project work She being an ideal person was always been my inspiration in every step
I offer my sincere thanks and gratitude to Professor KKBaruah Sir Tezpur University for his awful
lectures delivered in the regular classes which made me know my Project Topic more familiar before
head And along with his constant assistance suggestions guidance and co-operation and
encouragement during the entire course of project work
I would like to take the opportunity to thank and express my gratefulness to Dr KP Sarma Sir for
his guidance and providing me all the necessary fulfilments for the completion of the project work
I also convey my cordial thanks to DrRRHoque sir Head of the Department Department of
Environmental Science Tezpur University for giving me the golden opportunity to do this wonderful
project which not only helped me in doing a lot of Research but I explored many such events which
has been confined only to book and my limit of understanding was all based on theoretical
perspectives But after this project I could rather say that what I have gained is no more theoretical
now All I know is now practical
I also express and deliver my heartfelt Thanks to all the other Faculty members of Department of
Environmental Science- DrAKDas sir DrAshalata Devi marsquoam DrSSBhattacharya sir
4
DrMKumar sir Sumi Handique marsquoam for kind help and co-operation and encouragement during
the entire course of Project work
I would like to give my heartiest Thanks to Ms Bhaswatee Baroowa (Research Scholar) for being so
kind and helpful all throughout my Project She has been with me just like my elder sister in all phase
and makes me homely and giving moral support in every step
I express my cordial Thanks to my other Research Scholars who been giving constant advice
valuable suggestions and co-operation
Lastly but not least I would also like to thank my Parents My Local Guardian Dipankar Kalita
Department of Food Processing Technology Tezpur University and all my class-mate My Hostel
mates and my Roommate for being so humble and kind and along with helped me a lot in finishing
this project within the limited time
I am making this project not only for marks and partial fulfilment of the syllabus but its and handy
full aid to increase my knowledge and enlarge the circumference of my thoughts in every field
ldquoTHANKS AGAIN TO ALL WHO HELPED MErdquo
Abstract
Scarcity of water is a severe environmental constraint to plant productivity Drought-induced
loss in crop yield probably exceeds losses from all other causes since both the severity and
duration of the stress are critical Crop growth and productivity is adversely affected by
naturersquos wrath in the form of various biotic and abiotic stresses which adversely affects crop
growth and yield Therefore the present study was conducted with two major pulse crops
green gram (Vigna radiata L) and black gram (Vigna mungo L) in the sight to gain
information on various physiological changes taking place in the plants under low moisture
stress and its subsequent relation to plant growth development and yield Positive correlation
was found in plant height and leaf number and leaf area in both the pulses Regarding yield
both the pulses showed positive correlation with respect to soil moisture content The study
revealed that C1(T9) black gram is resistant than green gram against short-term moisture
stress And further it could be find out that the C3 (Pratap) of green gram has severe impact
on the moisture stress
5
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
DrMKumar sir Sumi Handique marsquoam for kind help and co-operation and encouragement during
the entire course of Project work
I would like to give my heartiest Thanks to Ms Bhaswatee Baroowa (Research Scholar) for being so
kind and helpful all throughout my Project She has been with me just like my elder sister in all phase
and makes me homely and giving moral support in every step
I express my cordial Thanks to my other Research Scholars who been giving constant advice
valuable suggestions and co-operation
Lastly but not least I would also like to thank my Parents My Local Guardian Dipankar Kalita
Department of Food Processing Technology Tezpur University and all my class-mate My Hostel
mates and my Roommate for being so humble and kind and along with helped me a lot in finishing
this project within the limited time
I am making this project not only for marks and partial fulfilment of the syllabus but its and handy
full aid to increase my knowledge and enlarge the circumference of my thoughts in every field
ldquoTHANKS AGAIN TO ALL WHO HELPED MErdquo
Abstract
Scarcity of water is a severe environmental constraint to plant productivity Drought-induced
loss in crop yield probably exceeds losses from all other causes since both the severity and
duration of the stress are critical Crop growth and productivity is adversely affected by
naturersquos wrath in the form of various biotic and abiotic stresses which adversely affects crop
growth and yield Therefore the present study was conducted with two major pulse crops
green gram (Vigna radiata L) and black gram (Vigna mungo L) in the sight to gain
information on various physiological changes taking place in the plants under low moisture
stress and its subsequent relation to plant growth development and yield Positive correlation
was found in plant height and leaf number and leaf area in both the pulses Regarding yield
both the pulses showed positive correlation with respect to soil moisture content The study
revealed that C1(T9) black gram is resistant than green gram against short-term moisture
stress And further it could be find out that the C3 (Pratap) of green gram has severe impact
on the moisture stress
5
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Keywords Crop growth Water deficiet Plant growth Morpho-physiological parameters
Yield responses
ContentsPage No
Chapter-1 Introduction 1-3
Chapter-2 Review of Literature 4-13
Chapter-3 Methods and Methodology 14-23
31 Experimental site 14-15
32 Preparation of soil beds 16
33 Experimental Design 16
34 Soil Analysis 16
341 Bulk density 17
342 Soil pH17
343 Soil Conductivity17
344 Moisture or Water content18
6
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
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in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
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Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
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and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
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Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
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Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
345 Soil Water holding Capacity19
346 Determination of available nitrogen in soil20-22
35 Plant Morphological Parameters22
36 Yield Parameters22-23
Chapter 4 Results and Discussions24-34
Chapter5 Conclusions35
Chapter6 References36-42
List of tables Page NoTable1 Physico-chemical parameters of soil 24Table2 Soil moisture content of Black gram 25Table3 Soil moisture content of Green gram 25Table4 Plant height of Black gram 26Table5 Plant height of Green gram26Table6 Leaf number of Black gram 27Table7 Leaf number of Green gram 27
7
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Table8 Leaf area of Black gram 28Table 9 Leaf area of Green gram28Table10 Pod number of Black gram28Table11 Pod number of Green gram29Table12 Yield in terms of GY DSI TDS MP RP of both the pulses29Table13 Crop yield of each variety under control and stressed condition29
List of figures Page NoFig 1 Preparation of soil beds14Fig 2 Study of Morphological Parameters14
8
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Fig 3 Experimental site at North Bank Plain Zone of Assam15Fig 4 Effect of low moisture stress on plant height of both the pulses30Fig 5 Effect of low moisture stress on leaf number of both the pulses 31Fig 6 Effect of low moisture stress on leaf area of both the pulses 32Fig 7(a) Effect of low moisture stress on Crop yield 33Fig 7(b) Effect of low moisture stress on Geometrical Yield 33Fig 8 (a) Effect of low moisture stress on mean productivity 34Fig8 (a) Effect of low moisture stress on rate productivity 34
9
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Chapter-1Introduction
lsquoIn a world that has the means for feeding its population the persistence of hunger is a
scandalrsquo (Food and Agriculture Organization 2006) This is a very comprehensive statement
made by the United Nations Organization and clearly explains the present state of apathy of
world agriculture While plant physiological investigations provide knowledge and better
understanding of the effects of abiotic stress on plants crop physiology provides information
on how stress affects plant growth and eventually agricultural yield Growing global demand
for food and feed of various plant-based products climatic change-imposed stress on
agriculture and the new opportunities such as the advances in molecular biology are covered
in this chapter Stress is an altered physiological condition caused by factors that tend to
disrupt the equilibrium Strain is any physical and chemical change produced by a stress
(Gaspar et al 2002) The term stress is used with various meanings the physiological
definition and appropriate term as responses in different situations The flexibility of normal
metabolism allows the response initiation to the environmental changes which fluctuate
regularly and are predictable over daily and seasonal cycles Thus every deviation of a factor
from its optimum does not necessarily result in stress Stress being a constraint or highly
unpredictable fluctuations imposed on regular metabolic patterns cause injury disease or
aberrant physiology The extent of crop yield loss due to abiotic stresses can be reduced by
manipulating plant metabolism and use of genetically-engineered plants This chapter
provides readers with a broad overview of the various types of abiotic stresses and their
influence on the yields of economically important crops such as cereals grain legumes
vegetable crops oilseed crops forage crops and medicinal crops (Rai and Takabe
2006)Abiotic stress such as drought flood problem soils (salinity and alkalinity) extreme
temperatures (especially at flowering and maturity times) chemical toxicity and oxidative
stress are serious threats to agriculture and the environment Increased salinity of arable land
is expected to have devastating global effects resulting in 30 land loss within the next 25
years and up to 50 by the year 2050 (Food and Agriculture Organization 2006) Stress in
10
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
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46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
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Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
biological terms means deviation in the normal physiology development and function of
plants which can be injurious and can inflict irreversible damage to the plant system
The type of stress that crop plants suffer from can be broadly grouped as lsquotemperature
variation at crucial stagesrsquo
There are several sticky abiotic parameters revolving around temperature eg frost damage
and evaporation stress Many living organisms parasitize plants such as the plant pathogenic
viruses bacteria fungi nematodes insects and phenerogamic plants Abiotic stresses are
inflicted by non-living things matter on which the plant system is dependent Ambient
temperature relative humidity sunshine microclimate soil nutrition soil biota and other
physico-chemical properties of the soil create stress on plant Stress is an altered
physiological condition caused by factors that tend to disrupt the equilibrium Strain is any
physical and chemical change produced by a stress (Gaspar et al 2002) The term stress is
used with various meanings the physiological definition and appropriate term as responses in
different situations The flexibility of normal metabolism allows the response initiation to the
environmental changes which fluctuate regularly and are predictable over daily and seasonal
cycles Thus every deviation of a factor from its optimum does not necessarily result in stress
Stress being a constraint or highly unpredictable fluctuations imposed on regular
metabolic patterns cause injury disease or aberrant physiology Plants are frequently exposed
to many stresses such as drought low temperature salt flooding heat oxidative stress and
heavy metal toxicity while growing in nature Drought is a meteorological term and is
commonly defined as a period without significant rainfall Generally drought stress occurs
when the available water in the soil is reduced and atmospheric conditions cause continuous
loss of water by transpiration or evaporation Drought stress tolerance is seen in almost all
plants but its extent varies from species to species and even within species
The severity of drought is unpredictable as it depends on many factors such as occurrence and distribution of rainfall evaporative demands and moisture storing capacity of soils (Wery et al 1994)
There are three types of drought-
Meteorological drought- Drought occurs when there is a prolonged period of below
average precipitation creating a natural shortage of available water
11
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
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48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
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Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Agricultural drought- Drought occurs when there isnrsquot enough moisture to support
average crop production on farms or average grass production on range land
Although agricultural drought often occurs during periods of average precipitation
when soil conditions or agricultural techniques require extra water
Hydrological drought- This type of drought occurs when water reserves in aquifers
lakes etc fall below an established statistical average
Of the various stresses known in nature drought stress poses a major threat to
crop production because water is essential at every stage of plant growth from seed
germination to plant maturation so degree of water is essential at every stage of plant
growthIts effect on growth and yield of different crops has been studied during the
last few years The drought environment has been reported to limit plant growth and
development prior to the loss of productivity especially of crop species
OBJECTIVES OF THE STUDY
The present investigation was undertaken with the following objectives
To see the effect of low moisture stress on the important morpho-physiological
parameters of two popular pulses black gram (Vigna mungo L) and green gram
(Vigna radiata L) cultivars commonly grown in Assam
To see how these parameters changes in the recovery period of low moisture stress
To find out the overall effect of low moisture stress on yield components of these
two cultivars of pulses
12
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Chapter-2Review of Literature
Of various abiotic stresses known in nature drought stress poses a major threat to crop
production because water is essential at every stage of plant growth from seed germination to
plant maturation (Chaves et al 2003 Athar and Ashraf 2005) so any degree of water
imbalance may produce deleterious effects on crop growth but it depends upon the nature of
crop species (El-Far and Allan 1995)Plantrsquos behaviour responses to drought are complex
and they adopted different mechanisms when encounter drought (Jones 2004) Upon
exposure to water deficit plants exhibit physiological biochemical and molecular responses
at both the cellular and whole plant levels (Greenway and Munns 1980 Hasegawa et al
2000) To cope with drought stress plants respond with morphological physiological and
biochemical changes These changes aim at the retention of water in spite of the high external
osmoticum and the maintenance of photosynthetic activity while stomatal opening is reduced
to counter water loss Long ago Turner (1979) described some mechanisms of water stress
tolerance in plants such as drought escape avoidance and tolerance to low water potential In
fact all these plant strategies depend on certain specific plant adaptations to water deficit
conditions (Turner 1979 1982 Chaves et al 2003) Under stress conditions plant cells
have the ability to prevent water loss and to maintain the continuous growth Plants
commonly react to these stresses by accumulation of compatible solutes such as proline in
cells which results in the improvement of environmental stress tolerance (Hong et al 2000
Ramajulu amp Sudhakar 2001 Ashraf and Foolad 2007) These solutes can be accumulated in
high concentrations without impairing plant metabolisms The accumulation of these
osmolytes may help plants to tolerate against stress by improving their ability to maintain
osmotic balance within the cell (Hasegawa et al 2000 Apse and Blumwald 2002) The
maintenance of turgor by osmotic adjustment is an importance of physiological adaptations
for minimizing the detrimental effects of water stress (Munns 2002)Water Deficiet Stress-
is one of the abiotic stresses that plants encounter between seedling to harvest stage When
soil moisture is continuously low water extraction by root and water transport within the
13
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
plant is reduced and a drought like situation prevails To overcome drought stress plants
respond by increasing the water extraction efficiency and the water use efficiency of roots
and simultaneously reduce the transpiration rate The first plant-stress symptom induced by
drought is often a rapid inhibition of shoot and to a lesser extent root growth (Hsiao 1973
Schulze 1986) In general shoot growth is more sensitive to water deficit than root growth
(Sharp and Davies 1989) The mechanisms underlying the sustained root growth under
water stress include osmotic adjustment (Saab 1992) and an increase in the loosening
capacity of the cell wall (Hsiao and Xu 2000) Plant growth is the result of daughter-cell
production by meristematic cell divisions and subsequent massive expansion of the young
cells Cell expansion is in turn dependent on biophysical changes which include a regulated
loosening of primary cell walls and subsequent yielding to the hydrostatic (turgor) pressure
generated by solute and water uptake into the cells (Neumann 1995 Cosgrove 1997) Cell
growth rate and its inhibition by water deficit are regulated by a complex multigenic series of
metabolic processes and are not simply a function of water availability for turgor
maintenance (Bassani et al 2004 Fan and Neumann 2004 Achard et al 2006 Fan et al
2006 Ma and Bohnert 2007 Smith and Stitt 2007)
Drought is the most important constraint to chickpea yield accounting for 40ndash50 yield
reduction globally (Chaves et al 2003 Athar amp Ashraf 2005) it was observed that four
approaches are being pursued by the chickpea plants to overcome the drought stress
bull High root mass
bull Smaller leaf area
bull Osmotic adjustment
bull Early-maturing short-duration varieties
bull Capacity of root to extract moisture from lower soil strata
bull Reduced evapo-transpiration
bull Elasticity of root system so that when soil cracks roots do not get clipped-off
bull Ideal maturity period
14
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
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Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
bull Self-created soil mulch
21Effects of drought stress on Plant Morpho- physiological Parameters
211 Effect on Plant height
In soybean the stem length was decreased under water deficit conditions (Specht et al
2001) The plant height was reduced up to 25 in water stressed citrus seedlings (Wu et al
2008) Stem length was significantly affected under water stress in potato (Heuer amp Nadler
1995) Abelmoschus esculentus (Sankar et al 2007 amp 08) Vigna unguiculata (Manivannan
et al 2007a) soybean (Zhang et al 2004) and parsley (Petroselinum crispum) (Petropoulos
et al 2008) Water stress greatly suppresses cell expansion and cell growth due to the low
turgor pressure
Osmotic regulation can enable the maintenance of cell turgor for survival or to assist plant
growth under severe drought conditions in pearl millet (Shao et al 2008) The reduction in
plant height was associated with a decline in the cell enlargement and more associated with
leaf senescence in A esculentus under water stress (Bhatt and Srinivasa Rao 2005) Water
stress reduced the head diameter 100- achene weight and yield per plant in sunflower There
was a negative correlation of head diameter with fresh root and shoot weight while a positive
one between dry shoot weight and achene yield per plant under water stress (Tahir and
Mehid 2001) Water stress for longer than 12 days at grain filling and flowering stage of
sunflower (grown in sandy loam soil) was the most damaging in reducing the achene yield in
sunflower (Mozaffari et al 1996 Reddy et al 2004) seed yield in common bean and green
gram (Webber et al 2006) maize (Monneveux et al 2006) and Petroselinum crispum
(Petropoulos et al 2008)Plant growth is greatly affected by water deficit At a
morphological level the shoot and root are the most affected and both are the key
components of plant adaptation to drought Plants generally limit the number and area of
leaves in response to drought stress just to cut down the water budget at the cost of yield loss
(Schuppler et al 1998) Greater plant fresh and dry weights under water limited conditions
are desirable characters
15
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
A common adverse effect of water stress on crop plants is the reduction in fresh and dry
biomass production (Farooq et al 2009) Plant productivity under drought stress is strongly
related to the processes of dry matter partitioning and temporal biomass distribution (Kage et
al 2004) Diminished biomass due to water stress was observed in almost all genotypes of
sunflower (Tahir and Mehid 2001)
Mild water stress affected the shoot dry weight while shoot dry weight was greater than root
dry weight loss under severe stress in sugar beet genotypes (Mohammadian et al 2005)
Reduced biomass was seen in water stressed soybean (Specht et al 2001) Poncirus
trifoliatae seedlings (Wu et al 2008) common bean and green gram (Webber et al 2006)
and Petroselinum crispum (Petropoulos et al 2008) A moderate stress tolerance in terms of
shoot dry mass plants was noticed in rice (Lafitte et al 2007)Impaired mitosis cell
elongation and expansion result in reduced plant height growth under drought (Nonami
1998 Kaya et al 2006 Hussain et al 2008)Water stress has been found to reduce greatly
the linear growth of shoots (Mazhar et al 1996 Mitchel et a 1997)
212 Effect on leaf area
Water stress induced reductions in growth and yield of several grain legumes are associated
with reductions in leaf area (Pandey et al 1984) Reduction in the average leaf size during
development occurs because later produced leaves are smaller while the older leaves that
abscise are larger Since leaf abscission increases under water stress a more uniform leaf size
during crop development with reductions in leaf area due to smaller average leaf size It is a
common observation that when server water deficiet develops lower (older) leaves are
desiccated and die first so as to reduce leaf area and water requirement than upper (younger)
leaves (Lopez et al 1996)Water deficit stress mostly reduced leaf growth and in turn the
leaf areas in many species of plant like Populus (Wullschleger et al 2005) soybean (Zhang
et al 2004) and many other species (Farooq et al 2009) Significant inter-specific
differences between two sympatric Populus species were found in total number of leaves
total leaf area and total leaf biomass under drought stress (Wullschleger et al 2005) The leaf
growth was more sensitive to water stress in wheat than in maize (Sacks et al 1997) Vigna
unguiculata (Manivannan et al 2007a) and sunflower (Manivannan et al 2007b amp 2008)
In the faba bean the lower leaf area under water stress is largely due to reduced leaf
expansion with relatively minor effects on leaf area production and death ( Kara Manos
16
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
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Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
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in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
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Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
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and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
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Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
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Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
1978 Farah 1981)For many grain legumes leaf area development (leaf production and
expansion) is more sensitive to water stress than leaf abscission ( Muchow et al 1985)
Vegetative growth of soyabean mild water stress reduces leaf expansion to a greater extent
than leaf production with little apparent affect on leaf senescence Severe water stress
however reduces leaf area largely by accelerated leaf senescence (Muchow et al 1986)
A number of studies have shown significant interaction between genotypes and water levels
(Siddique etal2000 Atteya et al 2003) Siddique et al (2000) studied response of four
cultivar of wheat to the four water levels It was shown that performance of these cultivars
was not similar over water level Atteya et al (2003) also indicated that exposure of plants to
drought led to noticeable decrease in leaf water potential relative water content and osmotic
potential However rate of dry matter accumulation was much lower in drought sensitive
genotypes as compared to the drought tolerant genotypes This may be due to higher
accumulation of cytokinins under drought stress Rauf and Sadaqat (2007) showed significant
higher accumulation of cytokinins in drought tolerant genotypes of sunflower They also
showed significant effect of endogenous plant growth regulators on the dry matter
partitioning It has been established that drought stress is a very important limiting factor at
the initial phase of plant growth and establishment It affects both elongation and expansion
growth (Anjum et al 2003a Bhatt amp Srinivasa Rao 2005 Kusaka et al 2005 Shao et al
2008) It has long been established that plants bearing small leaves are typical of xeric
environments Such plants withstand drought very well albeit their growth rate and biomass
are relatively low (Ball et al 1994) Leaf pubescence is a xeromorphic trait that helps protect
the leaves from excessive heat load Hairy leaves have reduced leaf temperatures and
transpiration (Sandquist and Ehleringer 2003) whilst inter- and intra-specific variation exists
for the presence of this trait Under high temperature and radiation stress increases the light
reflectance and minimizes water loss by increasing the boundary layer resistance to water
vapour movement away from the leaf surface Development of optimal leaf area is important
to photosynthesis and dry matter yield Water stress reduces seed yield in soybean usually as
a result of fewer pods and seeds per unit area (Specht et al 2001) In water stressed soybean
the seed yield was far below when compared to well-watered control plants (Specht et al
2001)Reducing the water content of plant resulted in a similar retardation in the time of leaf
emergence on the plants decline in their rate of growth and in their number (Shahid et al
1998)Various studies have indicated that water stress causes an inhibiting effect on the leaf
area of plants (Nielson et al 1998 Kawakami et al 2006 Guerrier et al 1990)
Muthechelian et al Franko et al (2006) have found direct relationships between each of
17
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
these growth parameters and the increasing rate of water stress Among the crops rice as a
submerged crop is probably more susceptible to drought stress than most other plant species
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006)
Under mild water stress situations leaf rolling reduces transpiration loss increases humidity
within the rolled leaf and thereby enables plants to withstand the prevailing harsh water
deficit stress conditions (Agarwal et al 2006)
213Effect on Root growth
In many succulents such as tomato potato celery etc the shallow spreading root system
quickly absorbs the small amount of water supplied by rain These ldquorain rootsrdquo develop
within few hours of shower mop-up the soil moisture and soon these roots dry out These
crops have an amazing ability to produce water absorbing fresh roots at a very high speed and
enable plants to overcome the water stress The aerial roots also absorb water from the
atmosphere under saturated relative humidity conditions or when there is a fine drizzle The
aerial roots of vanilla and orchids absorb water directly when misty conditions prevail
However the aerial roots produced by sorghum at the basal internodes are for reducing the
anaerobic stress under flooded conditions and serve as ldquobreathing rootsrdquo (Hasegawa et al 2000) Production of ramified root system under drought is important to above ground
dry mass and the plant species or decreased under pre-anthesis drought stress treatment in
wheat (Edward amp Wright 2008)Root profile and depth of plants change so to survive under
water deficit conditions Under upland conditions rice maize and sorghum have similar root
length density and water extraction patterns down to 60 cm depth (Levitt 1972) Below this
level water extraction by rice is insignificant and this is one of the reasons why rice is more
vulnerable to water stress compared to other crops Generally rice varieties with high root
length density tend to have high leaf water potential and delayed leaf senescence under water
deficit conditions Invariably rice cultivar with superior root length performs better under
water limiting conditions (Penna 2003 Reddy et al 2004 Agarwal et al 2006)Where repeated spells of transient drought occurs genotypes with larger root systems
tend to perform better Access to moist soil zone is accomplished by plants with maximum
rooting depth Species that have the capacity to establish roots in these lower moist soil layers
18
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
survive and contribute to plant population and community dynamics Since roots are the only
source to acquire water from soil the root growth its density proliferation and size are key
responses of plants to drought stress (Kavar et al 2007)
214 Effect on Plant reproductive Status
Male organelle development during meiosis in the microspore mother cells is extremely
vulnerable to water deficit conditions Water deficit adversely affects the development of
microspores or pollen grains causing pollen sterility Female fertility in contrast is relatively
unaffected by water stress at this stage The injury inflicted is apparently not by desiccation
of the reproductive tissue but as a consequence of water deficit in stems and leaves (Saini
1997) Stress-induced arrest of male gametophyte development is preceded by disturbances in
carbohydrate metabolism and distribution within anther An inhibition of the key sugar-
cleaving enzyme acid invertase decreases sugar delivery to reproductive tissues upon
inhibition of photosynthesis under stress conditions This is the signal that triggers metabolic
lesions leading to failure of male gametophyte development (Saini 1997)
215Effect on Fruit Development
In perennial fruit crops reduced moisture availability at flowering and fruit development
stages curtails pollen fertility berry formation and at subsequent stages causes fruit shedding
(Lindhauer et al 2007) Even fruits that eventually mature are of poor quality size and
appearance For example Vines nuts and melons produce unfilled fruits and nuts when
moisture stress occurs Early season water stress reduces the number of green leaves their
size shape and number of fresh leaves per flush In a row if annual cyclic water stress
occurs then it badly affects the canopy structure frame development number of branches
and the strength of twigs and woods
22 EFFECT ON PLANT YIELD COMPONENTS
The effects of water stress on growth and yield of different crops have been studied during
the last few years (Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga
et al 2003 Hussain et al 2004 Khan et al 2004 Rahman et al 2004) The crop species
19
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
show great differences for final harvestable yield under drought stress in early plantings of
sunflower the yield increase was associated with both an increase in grain number and in
individual grain weight (Soriano et al 2002)Exposure of sunflower plants to drought stress
at bud initiation stage was more detrimental to seed and biological yield than at seed filling
stage (Prabhudeva et al 1998)Giunta et al (1993) find that wheat yield decreased from 25 -
85 under drought stress
In some other studies on maize drought stress greatly reduced the grain yield which was
dependent on the level of defoliation due to water stress during early reproductive growth
(Kamara et al 2003 Monneveux et al 2006) Water stress reduces seed yield in soybean
usually as a result of fewer pods and seeds per unit area (Specht et al 2001) In water
stressed soybean the seed yield was far below when compared to well-watered control plants
(Specht et al 2001) There is an inadequacy of experimental evidence to summarises that
moisture stress in early stages affects the final yield of annual crops
With the normal weather thereafter loss is compensated by increased number of grains per
spike and better grain weight (DeLucia et al 1989) Lazaridou et al (2003)When
brief periods of moisture stress occurs the growth loss imposed on the plant is overcome
subsequently by increasing the functional efficiency Though the scars of stress will be
measurable on the crop productivity the magnitude of loss gets substantially reduced Plant
varieties possessing such alternate mechanisms to minimize the loss inflicted by drought are
rated as lsquotolerantrsquo
Three main mechanisms reduce crop yield by soil water deficit (i) reduced canopy
absorption of photosynthetically active radiation (ii) decreased radiation-use efficiency and
(iii) reduced harvest index (Earl and Davis 2003) Although plant responses to drought are
relatively well known plant performance under a more complex environment where multiple
stresses co-occur is fragmentary Hsiao and Xu al 2001) Drought-induced yield reduction
has been reported in many crop species which depends upon the severity and duration of the
stress period In maize water stress reduced yield by delaying silking thus increasing the
anthesis-to-silking interval This trait was highly correlated with grain yield specifically ear
and kernel number per plant (Cattivelli et al 2008) In pearl millet (Pennisetum glaucum)
co-mapping of the harvest index and panicle harvest index with grain yield revealed that
greater drought tolerance was achieved by greater partitioning of dry matter from stover to
grains (Yadav et al 2004)Under rainfed growing conditions seasonal fluctuations in water
availability may severely affect cereal grain yield during water deficient periods The most
suitable cultivar for such environments would produce high yields when rainfall is abundant
20
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
and without major yield reduction under rainfall shortages (Sinebo 2005)The extent of
variation in yield between stress and non stress conditions is widely accepted as a better
indicator of genotypic response to stress (Blum et al 1989)
Drought indices describing the relations between yield under stress and yield under
favourable conditions have been widely used (Golabadi et al 2006) These indices can be
used in the current study in identify potential tolerant andor high yielding cultivars suited to
drought prone growing conditions M Rashidil and K Seyfi (2007) conducted an experiment
on Cantaloupe (Cucumis melo) which is a very important vegetable crop of Iran to examine
the effect of drought stress on crop yield Water stress significantly affected crop yield of
cantaloupe in the order of 10 gt 30 gt 50 gt 70 available water deficit Highest crop
yield was obtained at 10 available water deficit and lowest value of crop yield was obtained
at 70 available water deficit treatment They concluded that higher value of crop yield
obtained at 10 available water deficit might be due to the more frequent application of
water resulting in more adequate moisture in active crop root zone sufficient moisture
conservation and better utilization of nutrients Lower crop yield obtained at 70 available
water deficit might be due to withdrawal of water application resulting in a lack of moisture
in active crop root zone inadequate moisture conservation and poor nutrient utilization
(Tahir amp Mehdi 2001 Aslam amp Tahir 2003 Ahmad et al 2003 Kumaga et al 2003
Hussain et al 2004 Khan et al 2004 Rahman et al 2004)Grain yield of single plant
decreases significantly with increasing severe drought stress and density Negative effect of
drought stress on yield component and grain in sunflower has been reported by other
researchers (Daulay et al 1983 Human et al 1998 Robinson et al 1985) Crop yield of
high osmotic-adjusting plants is more than that of low osmotic-adjusting plants Subbarao
Chauhan amp Johansen (2000) reported that osmotic adjustment was positively correlated with
grain yield It seems that the concentration and quality of cellular components is influenced
by severity of water deficit duration of recovery time and also by environmental conditions
such as different types of soil (Hoffman et al 2005 Ober et al 2005) It appears that the
drought stress-induced changes are reversible to some extent at least at the cellular level
Drought susceptibility index (S) represent drought tolerance at whole plant level regardless of
drought tolerance mechanism in operation (Grzesiak et al 1996 Vallejo and Kelly 1998)
The selected plants for lower drought susceptibility index may have diverse tolerance
mechanisms rather than based on single drought tolerant traits Therefore such type of
population may successfully cope with drought under range of environments Furthermore
drought tolerance is a complex phenomenon that does not always solely depend on single
21
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
plant trait Drought susceptibility index and fresh pod yield were independent of each other
and presence or absence of drought will establish their relationship (Falconer et al 1990)
In addition fresh pod yield of non-stress and drought condition should be considered as
separate resultant traits that were not always contributed by the same independent traits
(Falconer 1990) Similar type of relationship has also been obtained in other crops (Freres et
al 1989)Drought susceptibility of a genotype is often measured as a function of the
reduction in yield under drought stress (Blum 1988) whilst the values are confounded with
differential yield potential of genotypes (Ramirez and Kelly 1998) Rosielle and Hamblin
(1981) defined stress tolerance (TOL) as the differences in yield between the stress (Ys) and
non-stress (Yp) environments and mean productivity (MP) as the average yield of Ys and Yp
Fischer and Maurer (1978) proposed a stress susceptibility index (SSI) of the cultivar
Fernandez (1992) defined a new advanced index (STI= stress tolerance index) which can be
used to identify genotypes that produce high yield under both stress and non-stress
conditions Other yield based estimates of drought resistance are geometric mean (GM)
mean productivity (MP) and TOL The geometric mean is often used by breeders interested
in relative performance since drought stress can vary in severity in field environment over
years (Ramirez and Kelly 1998)Based on several studies (JLMaclagan 1993) proposed
that relative leaf water content can use as an index for rate of stress and wilting (Ritchie S
and T Henry 1990) in a study on two varieties of wheat used the relative leaf water content
as a stress index( Sullivan and JD Eastin 1994) proposed amount of electrolyte seepage in
plant leaves under stress condition (Winslow MD N Smirnoff 1984) showed that cell
membrane damage on plant which was under stress condition was lower than other
(RAFisher and R Maurer 1978) proposed stress susceptibility index to determine drought
tolerant variety Lower stress susceptibility index indicated higher toleration to drought The
selected varieties by this way have lower yield potential but in drought condition they will
produce higher yield (GCJ Fernandez 1993) proposed another index as a drought tolerance
index to determine drought tolerant variety Higher drought tolerance index show more
toleration to drought The variety that show higher drought tolerance index will produce
higher yield in stress and non stress condition (WA Iljin 1957)Among the stress
tolerance indicators a larger value of TOL and SSI represent relatively more sensitivity to
stress thus a smaller value of TOL and SSI are favoured In spring wheat cultivars Guttieri
et al (2001) using SSI criterion suggested that SSI more than 1 indicating above-average
susceptibility and SSI less than 1 indicated below-average susceptibility to drought stress
Ramirez and Kelly (1998) reported that GM and SSI as the mathematical derivations of the
22
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
same yield data selection based on a combination of both indices may provide a more
desirable criterion for improving drought resistance in common bean
Chapter-3Methods and methodologies
The experiment was carried out to evaluate the effect of short term moisture stress on some
important morpho-physiological parameters and yield components of the two popular pulses
black gram(Vigna mungoL) and green gram (Vigna radiataL) which are commonly grown in
Assam
31Description of the experimental site-
The present experiment was carried out during the period of August to December 2011 at
Tezpur University Campus which is located in North bank plain zone of Assam (26o14ʹ) and
(92o50ʹ)The soil of the experimental site is characterised by sandy loam textured having
slightly acidic in nature The soil physicochemical properties of the experimental site was
evaluated and presented in table no1 as shown in the chapter 4 The maximum and minimum
average temperature recorded during the experimental period ranges from 205o C to 328o C
and the average rainfall recorded was 0248mmAnd the relative humidity is 50 to 80
23
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
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Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
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Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
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in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
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Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
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and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
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Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
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Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Fig 1 Preparation of soil beds Fig 2 Study of Morphological Parameters
24
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Fig3 Experimental site at North Bank Plain Zone of Assam
25
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
32Preparation of soil beds
The experimental site was ploughed with the help of a tractor Manures were added and the
site was left for a week time so that the weeds and other microbes could be removed
Fertilizers like urea and SSP were applied 32 KGha and 220kgha respectively
Temporary polythene shed was constructed to create a temporary water stress condition
Removing of weeds and proper mixing of manure was followed by bed preparation A total
of 24 beds were prepared consisting of 9 rows and 4 columns maintaining requisite gaps
between plants and rows (P-P = 10Cm R-R =30Cm)
33 Experimental design
The experiment was conducted in Randomized block design (RBD) The varieties taken for
Black gram were C1-T9 C2-PU-39 and for green gram were C3-Pratap C4-TMB-37which
were collected from Regional Agricultural Research Station (RARS) Shillongoni Nagaon
We arranged 4 treatments -
C1 C2 C3 C4 - Were the control plants where watering was done regularly
C1T C2T C3T C4Tndash Were the treatment plants where watering was withdrawn for three
weeks during the reproductive stage We also arranged three replications for each treatment
Seeds were soaked in running tap water and then sown in the field on 4 th of September 2011
Watering was done in all the plots till 37 days And it was stopped in the plants which were
under treatment And thereafter again watering was done from the 58th day which starts with
the recovery period
The physico-chemical characteristics of soil were determined in the beginning of the experiment Change in morphological and biochemical characters were recorded during the stressed and the recovery period Finally the yield was also recorded
34 Soil Analysis
Soil Samples were collected in polythene bags and transferred into laboratory Then the
collected soils were dried and processed for further analysis by sieving through 2mm sieve
26
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
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Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
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Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
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Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Blum A 1996 Constitutive traits affecting plant performance under stress In
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Developing Drought and Low N Tolerant Maize pp 131ndash35
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Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
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Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
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Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
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Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
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Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
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Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
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Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
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Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
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Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
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Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
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Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
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Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
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Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
341 Bulk Density
The bulk density was determined by using the disturbed method In this method the weight of
an empty bottle without the stopper was first taken Then the bottle was filled with the dried
soil sample The bottle was tapped 15-20 times on a table by letting it fall from a height of
about 5 cm This tapping was assumed to produce the same intensity of packing of soil as
naturally occurring in the field Then the bottle was weighed again After that the bottle was
emptied and the volume of the bottle was measured by adding water from a burette up to the
rim
Bulk density was calculated as follows
W2 ndash W1
Bulk density of soil Db = -----------------------
V
Where
W1= weight of empty bottle
W2= weight of the bottle with soil
V = volume of water required to fill the bottle
342 Soil Reactions or Soil pH
Soil pH is a measure of the ldquohydrogen ion activityrdquo and depends largely on relative amount of
the absorbed hydrogen ions Thus it is a good measure of acidity and alkalinity of soil water
suspensions and provides a good identification of the soil chemical nature
pH of soil suspension highly depends on the soil water ratio and increases with dilution
Procedure
10 g of soil sample was mixed with 25 ml water to prepare the soil solution The solution was
stirred with a glass rod at regular interval for about 1 hour After that the pH of the
suspension was recorded with the help of digital pH meter
27
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
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metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
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Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
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Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
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Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
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Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
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Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
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Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
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Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
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Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
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Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
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Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
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Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
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Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
343 Soil Conductivity
10gm of soil sample was mixed with 50ml water to prepare the soil solution The solution
was then put in a shaker for 30 minutes After that with a digital electrical conductivity meter
conductivity of soil is recorded
344 Moisture or Water Content of the Soil
Water which occupies together with air the non solid porosity of soil is considered to be the
prime regulator of physical and chemical processes as well as biological activities in the soil
Water is essential for photosynthesis in the maintenance of soil turgidity Water acts
primarily as solvent and nutrient carrier from soil to the plant and then within the plant
Gravimetric method for soil moisture measurement
This is the standard and most simple method for determining the amount of water in soil The
principle involved is to place the weighed soil sample in an oven at 105oC and to dry it to a
constant weight The weight difference is considered to be the amount of water present This
water is expressed as a per cent of dry soil
APPARATUS
1 Sampling auger
2 Moisture cans
3 Balance with weights
4 Hot air oven
5 Desiccator
PROCEDURE
1 The clean dry moisture can along with its lid was weighed
2 The soil samples were collected with sampling auger from the required depth and
immediately about 50-100 g of soil was put in the moisture can and closed with the
tight fitting lid
3 The can containing soil sample was weighed with precision
4 The can containing moist soil was put in an oven The lid was removed and the soil
was dried at 105oC for 24 hours
28
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
5 After 24 hours the oven was switched off the sample was cooled for some time and
then transferred to a desiccator for further cooling The weight of the can with dried
soil was taken
6 Step 5 was repeated until a constant weight of can plus dry soil was obtained
CALCULATION
Weight of the empty can = W1
Weight of can + moist soil = W2
Weight of can + dry soil = W3
Amount of water present = W2 ndash W3
Weight of oven dry soil = W3 ndash W1
Weight of water
Soil moisture content by weight () = --------------------------------- x 100
Weight of oven dry soil
W2 ndash W3
= --------------------- x 100
W3 ndash W1
345 Soil Water holding Capacity
The water holding capacity of soil is defined as the capacity of water to hold water in plant
available form This is an important characteristic of soil for plant growth Soil that can hold
a lot of water support more plant growth and are less susceptible to leaching losses of
nutrients and pesticides
Procedure
1 The crushed soil sample was dried in an oven at 105oC overnight
2 A filter paper was placed inside the perforated bottom of the air cooled soil
box
3 The box was weighed and filled with dry soil
4 The box was placed in a petridish of 10cm diameter containing water for about
12 hour so that water enters the box and saturates the soil
5 The box is taken out from water wiped and then weighed again
29
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
6 The water holding capacity of the soil sample is calculated by using the
following formula
Water holding capacity () = (W2-W0)-(W1-W0)
-----------------------
W1-W2
Where W0= Weight of the empty box
W1= Weight of the box with soil
W2= Weight of the box with saturated soil
346 DETERMINATION OF AVAILABLE NITROGEN IN SOIL
PRINCIPLE
A given amount of soil is treated with an excess of alkaline KMnO4 and distilled KMnO4 is
a mild oxidizing agent in an alkaline medium (NaOH) Organic matter present in soil is
oxidized by the nascent oxygen liberated by KMnO4 in presence of NaOH and thus NH3
released is distilled and absorbed in a known volume of standard acid (boric acid) which is
titrated with HCl
REAGENTS
032 KMnO 4 32 g of KMnO4 was dissolved in distilled water and the volume was
made up to 1 litre
25 NaOH 25 g of NaOH pellets were dissolved in distilled water and volume
make up was done to 1 litre
4 Boric acid 40 g of H3BO3 was dissolved in 1 litre of distilled water
001N HCl
Liquid paraffin wax
Mixed indicator 03 g of bromocresol green and 02 g of methyl red was dissolved in
400 ml of 90 ethanol The indicator colour will change from red in acid solution to
blue in alkaline solution
30
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
PROCEDURE
1 3 g soil sample was taken in a 250 ml digestion tube
2 30ml 032 KMnO4 and 1ml liquid paraffin were added to it
3 Then the digestion tube was fitted well in the distillation unit for distillation
4 30 ml of 4 boric acid along with a few drops of mixed indicator were taken in a
250ml conical flask
5 The flask was fitted to the ammonia exhaust pipe
6 30ml 25 NaOH was then added to the contents of the digestion tube in three splits
7 Then the contents of the tube were distilled for 9 minutes and the distilled ammonia
was collected in the boric acid solution in the conical flask
8 After distillation was complete the collected ammonia solution was titrated with
001N HCl to original pink colour
9 A blank was carried out without soil following the same procedure
10 The burette reading (BR) was recorded
The titration part was carried out manually using gravimetric titration while the distillation
part was carried out using Kjeldahl methodology in a Nitrogen Analyzer having the following
details
Make Pelican Equipments
Model Digester KEL PLUS KES 12 L
Distiller KEL PLUS Distyl EM
CALCULATION
Correction of BR BRsample - BRblank
14 times BR times N of acid
Available Nitrogen () = ------------------------------------------------ x 100
Sample Weight times 1000
31
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Available Nitrogen (Kgha) = N times 2 times 104
14 times BR times 001
= ------------------------- times 100 times 224 times 104
3 times 1000
= BR times 1045
35 Plant Morphological parameters
The following plant morphological parameters were studied was studied at seven days
interval
Plant height-It was measured by a ruler scale
Leaf number It was counted manually in the field
Leaf area It was done with the help of graph paper
Pod number It was also counted manually
bull Root and Shoot Biomass Plant fresh weight was taken immediately after coming to
laboratory and for dry weight it was kept in the oven overnight at the temperature
80oC
36 Yield Parameters
On maturity of the plants harvesting was done on 26-11-2011 that was 37th day The
following yield attributing parameters like number of pod per plant number of seeds per pod
weight of each pod and seeds per pod and finally the weight of seeds per plant of treatment
including the control was calculated
Taking above parameters various indexes have been calculated out-
1) Harvest index Pod dry weight
-------------------------------------------
Total biomass on dry wt basis
32
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
2) Drought susceptibility index (1- Yield drought)
---------------------------------
Yield controlDII
Where DII = (1-mean yield droughtmean yield control)
3) Drought tolerance index (Yield stress times Yield control)
-----------------------------------------
Mean yield control
4) Mean productivity Yield control+ Yield drought
-------------------------------------------
2
5) Rate productivity Yield drought
--------------------
Yield control
6) Geometrical yield Yield drought
radic ----------------------
Yield control
37 Statistical Analysis
For Analyze of Correlation SPSS 160 software was used
33
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
CHAPTER 4 RESULTS AND DISCUSSIONS
RESULTS
1) Soil Physico-chemical Parameters The soil samples were collected and analyzed to determine its various physico-
chemical properties such as bulk density pH water holding capacity and soil
available nitrogen the result of which are presented in the table below
Table 1 Physico-chemical parameters of the soil of the experimental field
Sl No Soil Parameters Results
1 Bulk density 08825gmcc
2 Soil pH 585at 314oC
3 Water holding capacity 4573
4 Soil conductivity 53micro Siemenscm
5 Soil Available Nitrogen 374mgkg
Moisture content of soil
The soil moisture contents () of all the plots were determined before the onsets of
the treatments during the stress period and at drought recovery stages The results obtained
are presented in table 2 and table 3
Table 2 Soil moisture content () during the stress period and recovery stages of black gram
34
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
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Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
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Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
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followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Days after Sowing
Soil moisture content ()C1 C1T C2 C2T
17 9211 9343 9101 968627 9049 9201 9004 925137 9466 9179 9013 936744 9322 7093 9042 778651 9686 5915 9604 501358 9404 3327 9012 308465 9503 5094 9094 530873 9453 8020 9564 7689
Table3 Soil moisture content () during the stress period and recovery stages of green gram
Days after Sowing
Soil moisture content ()C3 C3T C4 C4T
17 9503 9547 9234 909527 9312 9138 9028 932437 9013 9100 9499 920844 8958 7306 9244 706251 9298 5567 9028 602258 9521 3696 8997 352165 9025 5494 9514 619273 9024 8439 9241 7902
2) Plant Morphophysiological Parameters
Plant morpho-physiological parameters such as plant height leaf number and leaf area and
pod number were studied at the beginning of treatment during the treatment period as well as
at recovery stages for both the crops
21) Plant height
Plant height was studied for each cultivar during treatment period and at recovery stages of
both black gram and green gram Drought stress has been found to decline the linear growth
of shoots in both the cultivars as compared to those of untreated ones
Upon re-watering both black gram and green gram plants started to recover in terms of
height but did not reach the control level The results are presented in Table 4 and Table 5
Table 4 Effect of low moisture stress on Plant Height (cm) of black gram
Days after Plant Height (cm)
35
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Sowing C1 C1T C2 C2T17 1883 1916 1765 156627 5507 2559 3766 213337 5666 2933 5633 253344 6066 3644 6133 293351 6166 4065 6366 313358 6837 4609 6533 376665 6533 4833 6767 403373 6613 4916 6866 4166
Table 5 Effect of low moisture stress on Plant Height (cm) of green gram
Days after Sowing
Plant Height (cm)C3 C3T C4 C4T
17 2016 1901 1866 181627 4133 2466 4133 256637 7466 2811 5566 291644 7766 3266 5806 343351 8066 3566 6033 370558 8302 3666 6366 418365 8302 4044 6466 453373 8633 4266 6533 4683
22) Leaf number
Leaf number is also an important parameter which is affected by drought and hence it was
studied for both the control and treatment plants in during the stressed as well as in the
recovery period During treatment period the number of leaves was found to be lesser than
that of the untreated plants and again it started to increase in the recovery period It can be
said that it was fluctuating The result is presented in table 6 and table 7
Table 6 Effect of low moisture stress on Leaf number of black gram
36
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
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to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Days after Sowing
Leaf numberC1 C1T C2 C2T
17 4 6 6 527 12 15 17 1137 18 23 27 1244 23 26 31 1451 23 25 33 1258 26 25 35 965 27 23 36 773 27 21 36 5
Table 7 Effect of low moisture stress on leaf number of green gram
Days after Sowing
Leaf numberC3 C3T C4 C4T
17 6 6 5 527 17 9 13 1037 25 12 17 1444 31 16 18 1451 32 16 22 1458 34 12 27 1265 34 11 28 873 34 11 29 4
23) Leaf area
Leaf area was found to be increasing at a faster rate during the previous part of the treatment
period but later on there was no further increase in the leaf area However in the recovery
period a little increment was observed Results are presented in Table 8 and Table 9
Table 8 Effect of low moisture stress on leaf area (cm2) of black gram
Days after Sowing
Leaf area (cm2)C1 C1T C2 C2T
17 506 526 45 53727 1031 1185 665 80137 2209 1356 947 80144 2203 149 1472 106751 2398 1137 1979 113958 2612 899 2166 93665 2814 688 2041 71973 2742 418 2404 536
Table 9 Effect low moisture stress on leaf area (cm 2) of green gram
Days after Leaf area ( cm2)
37
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Sowing C3 C3T C4 C4T17 446 613 503 34227 665 1211 905 62537 947 1548 905 85944 1472 1437 1677 119251 1979 1408 2071 10558 2166 1045 2221 75865 2041 891 2471 50973 2404 647 2766 335
24) Pod number
Pod number for each cultivar was recorded and it was found that it increased at a faster rate at
the beginning but later due to moisture stress the increment in the number of pods fell down
and got reduced as compared to the control plants Table 10 and Table 11 show the effect of
moisture stress on the number of pods of both the control and stressed plants
Table 10 Effect of low moisture stress on pod number of black gram
Days after Sowing
Pod numberC1 C1T C2 C2T
17 - - - -27 - - - -37 7 3 4 444 8 5 5 351 11 7 9 458 12 8 9 465 15 7 12 573 17 6 15 4
Table 11 Effect of low moisture stress on pod number of green gram
Days after Sowing
Pod numberC3 C3T C4 C4T
17 - - - -27 - - - -37 5 3 5 244 5 5 6 351 10 6 7 358 13 6 14 465 15 6 14 573 14 4 13 3
3) Yield and Yield Attributing Parameters
38
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Drought stress was found to have significant effect on all the cultivars It severely
reduced crop yield of both the pulses The yield of both green gram and black gram were
calculated in terms of Gross yield (GI) Drought susceptibility index (DSI) Drought tolerance
index (DTI) Tolerance drought stress (TDS) Mean productivity (MP) and Rate productivity
(RP) The values of which are presented in table 12
Table12 Crop yield in terms of GY DSI DTI TDS MP and RP of both the pulses
Varieties GY DSI DTI TDS MP RP
C1 086 2026 077 -114 86021 -183
C2 084 1891 073 -171 40277 -581
C3 057 1054 034 076 15825 -1153
C4 085 666 075 -159 13485 -254
The crop yield on the basis of qha was also calculated out for both the control and treatment
plants of all four cultivars and these data are presented in table 16 below-
Table13 Crop yield of each variety under control and stressed condition
DISCUSSIONS
Water stress posses a major threat to crop production worldwide Water deficitdrought
affects every aspect of plant growth and the yield modifying the anatomy morphology
physiology biochemistry and finally the productivity of crop (Jones et al 2003 Hafiz et al)
because water is essential at every stage of plant growth from seed germination to plant
maturation (Chaves et al 2003) so any degree of water imbalance may produce deleterious
effects on crop growth (El-Far etal 1995) Yegappan et al (1982) found that drought stress
reduced number of leaf head diameter leaf area weight of 1000 grains and grain yield
significantly In the present study the plant height and the leaf area showed a slight decline
during the water stress period Upon rewatering both black gram and green gram plants
started to recover in terms of their growth
39
Treatments Crop Yield (qha) Treatments Crop Yield (qha)C1 2321 C3 1642
C1T 1729 C3T 477
C2 2182 C4 792
C2T 1609 C4T 538
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Plant Height
Water stress has been found to reduce greatly the linear growth of shoots (Mazhar et al
1996 Mitchel etal 1997) In the present study the plant height and the leaf area showed a
slight decline during the stress period which coincides with the results obtained by Nielson et
al (1998) Kawakami et al (2006) Guerrier et al (1990) Water stress had the drastic
effect on the plant height of all the wheat varieties in the experiment conducted by Ahmad et
al (2009) The wheat varieties used in this experiment showed significant differences
(plt005) between them an increasing level of water stress for plant height This reduction in
plant height is generally associated with a decline in the cell enlargement and more leaf
senescence under water stress as concluded by Bhatt amp Srinivasa Rao (2005)
Fig4 Effect of low moisture stress on plant height of both the pulses
From the analysis of variance (ANOVA) it was found that regarding plant height black gram
is significant as the calculated value was smaller than the tabulated value However green
gram was found insignificant as its calculated value was larger than the tabulated value On
finding out the correlation in SPSS 160 software it was found that plant height is positively
correlated with soil moisture content in both black gram and green gram
40
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Leaf number
After studying the leaf number of both black gram and green gram it was found that at initial
stage of treatment it was increasing but later due to less moisture availability defoliation was
seen more in green gram compared to black gram Similar results were obtained by Sacks et
al (1997) Manivannan et al (2007 amp 2008) while working with maize and sunflower
Water deficit stress mostly reduced leaf growth and in turn the leaf areas in many species of
plant like Populus (Wullschleger et al 2005) soybean (Zhang et al 2004) In some other
studies it was also concluded that drought stress greatly reduced the grain yield which was
dependent on the level of defoliation ie reduction in leaf number due to water stress during
early reproductive growth (Kamara et al 2003 Monneveux et al 2006) A major effect of
drought is reduction in photosynthesis which arises by a decrease in leaf expansion impaired
photosynthetic machinery premature leaf senescence and associated reduction in food
production (Wahid and Rasul 2005)
Fig 5 Effect of low moisture stress on plant leaf number of both the pulses
On analysing the variance it was found that black gram is significant compared to green gram
in terms of leaf number On finding out correlation between leaf number and soil moisture
content it was found positive for both the pulses
41
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Leaf area
Water stress has been found to reduce leaf area photosynthesis leaf chlorophyll contents and
consequently grain yield (Jun-Chen and Dai-Junying 1996)An increase in plant leaf area
resulted in an increase in dry matter production at a similar rate for both non-stress and stress
conditions Cultivar and water availability determined leaf area development and in turn
leaf area determined the amount of dry matter produced (Maria Gomez et al 2005) In our
experiment also leaf area has been found to be increasing before the onset of the treatment
During the initial stage of treatment it was increasing later on leaf expansion was reduced
However during recovery period it again showed a level of increase The reduced leaf area
observed in the stressed plants resulted primarily from a mitotic sensitivity to water stress in
cotton leaves (Jerry Berlin et al 2007) Leaf area distribution between primary and lateral
shoots was not significantly modified by water stress The capacity to form leaf area and the
sensitivity of leaf area development to water stress were apparently under genetic control
(Constantino Ruiz et al 2005)
Fig6 Effect of low moisture stress on plant leaf area of both the pulses
On analysing the variance it was again found that black gram is significant compared to green
gram in terms of leaf area On finding out correlation between leaf area and soil moisture
content it was found positive for both the pulses
42
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Yield and yield attributing parameters
Many yield-determining physiological processes in plants respond to water stress Yield
integrates many of these physiological processes in a complex way Thus it is difficult
to interpret how plants accumulate combine and display the ever-changing and indefinite
physiological processes over the entire life cycle of crops (Nonami 1998 Kaya et al 2006
Hussain et al 2008) Drought stress in soybean reduced total seed yield and the branch seed
yield (Frederick et al2001)Yield is generally correlated with the length of crop duration
under favourable growing conditions and any decline in crop duration below the optimum
would tax yield (Turner et al 2001)Water stress reduces seed yield in soybean usually as a
result of fewer pods and seeds per unit area (Specht et al 2001)Yield attributing parameters
such as crop yield geometrical yield drought tolerance index drought susceptibility index
tolerance drought stress mean productivity and rate production
(a) (b)
Fig7 Relationship of low moisture stresses on (a) crop yield and (b) geometrical yield of
black gram and green gram
43
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
(a) (b)
Fig 8 Relationship of low moisture stress on mean productivity and rate productivity
In the present experiment the biomass yields and seed yield of both the pulses (black gram
and green gram) were found to increase in control sets almost in all levels as found out by
(Cure and Acock 1986 Roggers and Dahlmann 1993) However impact was seen in the
treatment sets in all levels especially more impact was seen more in C3 variety of green gram
than both the varieties of black gram Similar results were also by (Daneshian J and P
Jonobi et al 2001) that drought stress is one of the important soybean growth limiting
factor which decreases plant growth during vegetative stage Chimenti et al (2002) Erdem
et al (2006) indicated that grain yield and weight of 1000 grains decreased with increasing
drought stress Karam et al (2007) showed that with increasing drought stress leaf area
index grain yield and its component decreased which is also found in our study
According to other yield attributing parameters like drought susceptibility index drought
tolerance index mean productivity and rate productivity the black gram variety T9 is more
resistant than PU-39 and for green gram Pratap is more resistant than TMB-37 against
drought stress which coincides with the findings of Azimzadeh et al (2011) and Toker et al
(1996) Positive correlation was obtained between yield of both the pulse cultivars and soil
moisture content with a value of 0415 in black gram and in green gram it is 061 Although
the applied drought significantly affected all the cultivars by drought stress leading to loss of
growth productivity and yield the impact was somewhat lesser in black gram cultivars in
comparison to green gram cultivars which reveal that black gram is more resistant than green
gram against drought stress
44
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Chapter5
ConclusionThe present experiment was conducted with the two common pulse crops namely black gram
(Vigna mungoL) and green gram (Vigna radiataL) with the objective to study the morpho-
physiological changes that took place in response to the low moisture stress The morpho-
physiological parameters such as plant height leaf number leaf area pod number were
studied under moisture stress condition as well as subsequent recovery stage
On maturation yields of these two crops were recorded The study revealed that moisture
stress has significant impact on all these parameters Plant height was seen to be increased in
the initial part of the treatment however it was reduced during the treatment period however
there was an increase in the height during the recovery period Positive correlation was seen
between plant height and soil moisture for both black gram and green gram Leaf number was
also seen to be reducing in number due to less availability of moisture in the soil But more
impact was seen in green gram In case of leaf area both black gram and green gram showed
positive correlation with the soil moisture content which can be inferred that leaf growth is
very much susceptible to moisture content Lastly it can be concluded that all the
morphological parameters had positive correlation with the soil moisture content And that if
we compare the impact of low moisture stress on both the pulse crops then green gram is
more sensitive compared to black gram which proofed that black gram C1(T9) is resistant to
low moisture stress and C3(Pratap) is susceptible to moisture stress
45
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Chapter-6References
Abbadi J and J Gerendas (2009) Nitrogen use efficiency of Safflower as compared
to sunflower J Pant Nutrition Vol 32(6) 929-945
Abdul Jaleel C Manivannan P Sankar B Kishorekumar A Gopi R
Somasundaram R Panneerselvam R (2007) Water deficit stress mitigation by
calcium chloride in Catharanthus roseus Effects on oxidative stress proline
metabolism and indole alkaloid accumulation Colloid Surf B 60 110ndash116
Agarwal PK Agarwal P Reddy MK Sopory SK (2006) Role of DREB
transcription factors in abiotic and biotic stress tolerance in plants Plant Cell Rep 25
1263ndash1274
Aharon R Shahak Y Wininger S Bendov R Kapulnik Y Galili G (2003)
Overexpression of a plasma membrane aquaporins in transgenic tobacco improves
plant vigour under favourable growth conditions but not under drought or salt stress
Plant Cell 15 439ndash447
Ahmad S H R Ahmad M Y Ashraf M Ashraf and E A Waraich (2009)
Sunflower (Helianthus annuus L) response to drought stress at germination and
seedling growth stages Pak J Bot 41(2) 647-654
Aless G G F Power and D C Zimmerman (1997) Sunflower yield and water use
as influenced by planting date populateion and row spacing Agron J 69 465-469
Ali Meo A (2000) Impact of variable drought stress and nitrogen levels on plant
height root length and grain numbers per plant in a sunflower (Helianthus annuus L)
Var Shams Pah J Agri Sci Vol 37(1-2) 89-92
Allison J C S and R J Haslam(1993) Theoritical assessment of potential for
increasing productivity of sugarcane throught increased nitrogen fertilization Proc
South African Sugar Technol Assoc 57-59
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003a Water stress in
barley (Hordeum vulgare L) I Effect on morphological characters Pakistan J
Agric Sci 40 43ndash44
46
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Anjum F M Yaseen E Rasul A Wahid and S Anjum 2003b Water stress in
barley (Hordeum vulgare L) II Effect on chemical composition and chlorophyll
contents Pakistan J Agric Sci 40 45ndash49
Ashraf MY AR Azmi AH Khan and SA Ala 1994 Effect of water stress on
total phenols peroxidase activity and chlorophyll content in wheat (Triticum aestivum
L) genotypes under soil water deficits Acta Physiol Plant 16 185ndash191
Begg J E and Turner N C 1976 Crop Water Deficit Adv in Agron 28 161-217
Bhatt RM and NK Srinivasa Rao 2005 Influence of pod load response of okra to
water stress Indian J Plant Physiol 10 54ndash59
Blum A 1983 Genetic and Physiological Relationships in Plant Breeding for
Drought Resistance Agr Water Manage 7 195-205
Blum A 1996 Constitutive traits affecting plant performance under stress In
Edmeades GO M Banziger HR Mickelson and CB Pena- Valdivia (eds)
Developing Drought and Low N Tolerant Maize pp 131ndash35
Burow M D and Coors J G 1994 Diallel A Microcomputer Program for the
Simulation and Analysis of Diallel Crosses Agron
Buschmann C Lichtenthaler HK 1998 Principles and characteristics of multi-colour
fluorescence imaging of plants Journal of Plant Physiology 152 297ndash314
Chaves MM JS Pereira J Maroco ML Rodriques CPP Ricardo ML Osorio
I Carvatho T Faria and C Pinheiro 2002 How plants cope with water stress in the
field photosynthesis and growth AnnBot 89 907ndash916
Chaves MM Oliveira MM 2004 Mechanisms underlying plant resilienceto water
deficits prospects for water-saving agriculture Journal of Experimental Botany 55
2365ndash2384
Farooq M A Wahid N Kobayashi D Fujita and SMA Basra 2009 Plant
drought stress effects mechanisms and management Agron Sustain Dev 29 185ndash
212
Farooq M SMA Basra A Wahid ZA Cheema MA Cheema and A Khaliq
2008 Physiological role of exogenously applied glycinebetaine in improving drought
tolerance of fine grain aromatic rice (Oryza sativa L) J Agron Crop Sci 194 325ndash
333
Fischer RA Maurer R Drought Resistance in Spring Wheat Cultivars I Grain
Yield Response Aust J Agric Res 29 897-912 1978
47
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Flenet F A Boundiols and C Suriava (1996) Sunflower response to a range of soil
water contents European Journal of Agronomy 15 161-167
Gardner F P R B Piers and R L Michel (1994) Agronomy plant physiology
Mashhad JDM press P 467 (Translation in Persian)
Gaspar T T Franck B Bisbis C Kevers L Jouve JF Hausman and J Dommes
2002 Concepts in plant stress physiology Application to plant tissue cultures Plant
Growth Regul 37263ndash285
Gubbels G H and W Dedio (1999) Effect of plant density and soil fertility on oil
seed sunflower genotypes Can J Pl Sci 66 (3)521-527
Harris HC Williams JR Mason MK (1978) Influence of temperature on oil content
and composition of sunflower seed Aust J Agric Res 291203ndash1212
Hasanzade A (2002) The effect of different amounts of Nitrogen fertilizer on yield
and yield component and grain oil of sunflower Uremia Agri SCI Research 2125-
33
Hashemi-Dezfouli A and S J Herbert (1992)Effect of leaf orientation and density
on yield of corn Iran Agric Res 11 89 - 104
Havaux M Strasser RJ Greppin H 1991 A theoretical and experimental analysis of
the qp and qnp coefficients of chlorophyll fluorescence quenching and their relation
to photochemical and nonphotochemical events Photosynthesis Research 27 41ndash55
Havaux M 1998 Carotenoids as membrane stabilizers in chloroplasts Trends Plant
Sci 3 147ndash151
Heuer B and A Nadler 1995 Growth development and yield of potatoes under
salinity and water deficit Australian J Agric Res 46 1477ndash1486
Human J J D Dutoit H D Benzuid Enhout and L P Bruyn (1998) The
influence of plant water stress on net photosynthesis and yield of
SunflowerAgriculttural University of south Africa Crop Science 164(4)231-241
Hura T Grzesiak S Hura K Grzesiak MT Rzepka A 2006Differences in the
physiological state between triticale and maize plants during drought stress and
followed rehydration expressed by the leaf gas exchange and spectrofluorimetric
methods Acta Physiologiae Plantarum 28 433ndash443
Jaleel CA B Sankar PV Murali M Gomathinayagam GMALakshmanan and
R Panneerselvam 2008e Water deficit stress effects on reactive oxygen metabolism
48
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
in Catharanthus roseus impacts on ajmalicine accumulation Colloids Surf B
Biointerfaces62 105ndash111
Jaleel CA P Manivannan A Kishorekumar B Sankar R Gopi R Somasundaram
and R Panneerselvam 2007c Alterations in osmoregulation antioxidant enzymes
and indole alkaloid levels in Catharanthus roseus exposed to water deficit Colloids
Surf B Biointerfaces 59 150ndash157
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007a Pseudomonas fluorescens enhances biomass yield and
ajmalicine production in Catharanthus roseus under water deficit stress Colloids
Surf B Biointerfaces 60 7ndash11
Jaleel CA P Manivannan B Sankar A Kishorekumar R Gopi R Somasundaram
and R Panneerselvam 2007b Water deficit stress mitigation by calcium chloride in
Catharanthus roseus effects on oxidative stress proline metabolism and indole
alkaloid accumulation Colloids Surf B Biointerfaces 60 110ndash116
Johansen B Baldev B Brouwer JB Erskine W Jermyn WA Li-Juan L
Malik BA Ahad Miah A Slim SN Biotic and Abiotic Stresses Constraining
Productivity of Cool Season Food Legumes in Asia Africa and Oceania in
Expanding the Production and Use of Cool Season Food Legumes Eds FJ
Muehlbauer and WJ Kaiser Kluwer Academic Pub printed theNetherlands p175-
194 1994
Kalra N Aggarwal PK Chander S Pathak H Choudhary R Chaudhary A Mukesh S
Rai HK Soni UA Anil S Jolly M Singh UK Owrs A Hussain MZ (2003) Impacts
of climate change on agriculture In Shukla PR Subodh KS Ravindranath NH Garg
A Bhattacharya S (eds) Climate change and India vulnerability assessment and
adaptation University Press India pp 191ndash226
Kozlowski TT (1997) Responses to woody plants to flooding and salinity Tree
physiology Monograph No 1 Heron Publishing Victoria Canada
Levitt J (1972) Responses of plants to environmental stresses Academic Press New
YorkSan FranciscoLondon
Lopez MT Toojinda T Vanavichit A Tragoonrung S (2003) Microsatellite markers
flanking the tms2 gene facilitated tropical TGMS rice line development Crop Sci
432267ndash2271
Majee M Maitra S Dastidan GK Patnaik S Chatterjee A Hait NC Das KP
Majumdar AL (2004) A novel salt tolerant L-myo-Insositol-1-phosphate synthase
49
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
from Proteresia Coarctata (Roxb) Tateoka a haplophytic wild rice J Biol Chem
2728539ndash28552
Matthews MA Boyer JS 1984 Acclimation of photosynthesis to low water
potentials Plant Physiology 74 161ndash166
Mohammadian R M Moghaddam H Rahimian and SY Sadeghian 2005 Effect
of early season drought stress on growth characteristics of sugar beet genotypes
Turkisk J Bot 29 357ndash368
Monneveux P C Saacutenchez D Beck and GO Edmeades 2006 Drought tolerance
improvement in tropical maize source populations evidence of progress Crop Sci
46 180ndash191
Mozaffari K Y Arshi and H Zeinali-Khanghaa 1996 Research on the effects of
water stress on some morphophysiological traits and yield components of sunflower
(Helianthus annuus L) Seed Plant 12 24ndash33 Nam NH YS Chauhan and C
Johansen 2001 Effect of timing of drought stress on growth and grain yield of extra-
short-duration pigeonpea lines J Agric Sci 136 179ndash189
Nayyar H S Kaur S Singh and HD Upadhyaya 2006 Differential sensitivity of
Desi (small-seeded) and Kabuli (large seeded) chickpea genotypes to water stress
during seed filling effects on accumulation of seed reserves and yield J Sci Food
Agric 862076ndash2082
Petropoulos SA Dimitra Daferera MG Polissiou and HC Passam 2008 The
effect of water deficit stress on the growth yield and composition of essential oils of
parsley Sci Hort 115 393ndash397
Prabhudeva TV MM Chalapathi S Thimmegowda N Devakhumar GG Rao
and K Mallikarjuna 1998 Soil moisture stress and drought susceptibility index in
sunflower Indian Agric 42 287ndash289
Prochazkova D RK Sairam GC Srivastava and DV Singh 2001 Oxidative
stress and antioxidant activity as the basis of senescence in maize leaves Plant Sci
161 765ndash777
Razmjoo K P Heydarizadeh and MR Sabzalian 2008 Effect of salinity and
drought stresses on growth parameters and essential oil content of Matricaria
chamomile Int J Agric Biol 10 451ndash454
Reddy AR KV Chaitanya and M Vivekanandan 2004 Drought induced
responses of photosynthesis and antioxidant metabolism in higher plants J Plant
Physiol 161 1189ndash1202
50
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Sacks MM WK Silk and P Burman 1997 Effect of water stress on cortical cell
division rates within the apical meristem of primary roots of maize Plant Physiol
114 519ndash527
Sadiq II Siddiqui KA Arain CR Azmi AR Wheat Breeding in a Water-
Stress Environment I Delineation of Drought Tolerance and susceptibility Plant
Breeding 113 36-46 1994
Samarah NH 2005 Effects of drought stress on growth and yield of barley Agron
Sustain Dev 25 145ndash149
Saxena MC The Challenge of Developing Biotic and Abiotic Stress Resistance in
Cool-Season Food Legumes in Breeding for Stress Tolerance in Cool-Season Food
Legumes Edited by KB Singh and MC Saxsena A Wiley-Sayce Publication p3-
14 1993
Sevilla Spain JA Consejeriacutea de Agricultura y Pesca Specht JE K Chase M
Macrander GL Graef J Chung JP Markwell M Germann JH Orf and KG
Lark 2001 Soybean response to water A QTL analysis of drought tolerance Crop
Sci 41 493ndash509
Singh KB Malhotra RS Halila MH Knights EJ Verma MM Current Status
and Future Strategy in Breeding Chickpea for Resistance to Biotic and Abiotic
Stresses in Expending the Production and Use of Cool Season Food Legumes Eds
FJ Muehlbauer and WJ Kaiser Klwer Academic Pub printed the Netherlands p
572-591 1994
Singh V S K Sharma BI Verma and V Singh (1995) Response of rainy season
sunflower (Helianthus annuus) to irrigation and nitrogen under north western
Rajasthan Indian Journal of Agronomy 40 239-242
Smirnoff N 1993 The role of active oxygen in the response of plants to water
deficit and desiccation New Phytol 125 27ndash58
Soriano MA FJ Villalobos and E Fereres 2002 Stress timing effects on
sunflower harvest index In Villalobos FJ and L Testi (Eds) VII Congress of the
European Society for Agronomy pp 141ndash2
Taghdiri B G Ahmadvan and H A Mazaheri Laghab (2006) The effect of plant
spacing on yield and yield components of four sunflower cultivars Agri Res 6(1)
26-35
51
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52
Tahir MHN and SS Mehid 2001 Evaluation of open pollinated sunflower
(Helianthus annuus L) populations under water stress and normal conditions Int J
Agric Biol 3 236ndash238
Tahir MHN M Imran and MK Hussain 2002 Evaluation of sunflower
(Helianthus annuus L) inbred lines for drought tolerance Int J Agric Biol 3 398ndash
400
Tahkokorpi M K Taulavuori K Laine and E Taulavuori 2007 After effects of
drought-related winter stress in previous and current year stems of Vaccinium
myrtillus L Environ Exp Bot 61 85ndash93
Wery J Adaptation to Frost and Drought Stress in Chickpea and Implications in
Plant Breeding in MC Saxena JI Cubero and J Wery (eds) Present Status and
Future Prospect of Chickpea Crop Production and Improvement in the Mediterranean
Countries Options Mediterraneennes Serie A Seminaires Mediterranees No9
Zarogosa Spain CIHEAM p77-85 1990
Wullschleger SD TM Yin SP DiFazio TJ Tschaplinski LE Gunter MF
Davis and GA Tuskan 2005 Phenotypic variation in growth and biomass
distribution for two advanced-generation pedigrees of hybrid poplar Canadian J For
Res 35 1779ndash1789
Zaffaroni E and A A Schneiter (1991) Sunflower production as influenced by plant
type plant population and row arrangement Agron J 63113-118
Zhang M L Duan Z Zhai J Li X Tian B Wang Z He and Z Li 2004 Effects
of plant growth regulators on water deficit-induced yield loss in soybean Proceedings
of the 4th International Crop Science Congress Brisbane Australia
Zhao CX LY Guo CA Jaleel HB Shao and HB Yang 2008 Prospects for
dissecting plant-adaptive molecular mechanisms to improve wheat cultivars in
drought environments Comp Rend Biol 331 579ndash586
52