topics term paper assignments¹€อกสาร... · 1 reducing water use, increasing yields...
TRANSCRIPT
1
123752 Horticultural Crops Breeding for Pest
Resistance and Environmental Tolerance
123752: Suchila, KKU, Thailand
Assoc. Prof. Suchila Techawongstien (D.Agr.) Plant Breeding Research Center for Sustainable
Agriculture Faculty of Agriculture, Khon Kaen University
1
Course description
Relationship between plant genetics, diseases, insects and environmental stress; Techniques of crop improvement for
123752: Suchila, KKU, Thailand
resistance to diseases, insects and environmental stress; and
Management of crop resistant variety
2
Topics
Chapter I: Introduction (Terminology, Importance of resistant variety and Principles and methods of breeding for resistance): slide 5‐23
Chapter II: Inheritance of resistance/tolerance traits (Gene action and gene interaction, Sources of resistance, Heritability and breeding method, and assessment of resistance/tolerance): slide
123752: Suchila, KKU, Thailand
24 ‐158
Chapter V: Plant and environment relationship (Stress environment, Responses of plant to environmental stress and Plant breeding and yield stability): slide 159‐192
Chapter VI: Management of resistance/tolerance varieties: slide 193‐199 3
Term paper assignments
Search and review the update papers Present and discuss the update
papers Suggested topics: bacteria, fungi,
123752: Suchila, KKU, Thailand
Suggested topics: bacteria, fungi, virus, nematode, insect-pests and environmental resistance/tolerance of economic plants (or the major crops of the student’s research)
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Lecture Chapter I: Introduction
1.1 Terminology
1 2 Importance of resistant variety
Chapter I: Introduction
123752: Suchila, KKU, Thailand
1.2 Importance of resistant variety
1.3 Principles and methods of
breeding for resistance
5
1.1 Terminology
1) Parasite
2) Pathogen
3) Disease
7) Tolerance
8) Disease escape
9) Immunity
) ff l h
1.1 Terminology
123752: Suchila, KKU, Thailand
3) Disease
4) Resistance
5) Susceptible
6) Hypersensitivity
10) Differential host
11) Physiological
race
12) Pathogenicity
6
1
1.1 Terminology (cont.)
13) Vertical resistance14) Horizontal resistance
18) Preference19) Non‐ preference20) Environmental stress
Chapter I: Introduction
123752: Suchila, KKU, Thailand
15) Disease assessment 16) Persistent vector17) Non‐ persistent vector
21) Phenotypic responses22) Genotypic responses 23) Stability
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Importance of resistant variety
In 1845-46: Irish Famine-potato late blight (Phytophthora infestans)
40% of potato tuber in Belgium/Ireland was rotten by late blight
1. 2 Importance
123752: Suchila, KKU, Thailand
More than 1.5 million people died (Population of Ireland declined from 8.2 to 6.2).
https://www.google.co.th/search?q=potato+late+blight+pictures8
At Bengal: Epidemic of Brown Spot (Helminthosporium oryzae) in rice
75-90% of rice yield in Bengal was decreased
Importance of resistant variety (cont.)
1. 2 Importance
123752: Suchila, KKU, Thailand
More than 2 million people died
Europe: Epidemic of corn caused >20% of maize yield
9
Hurricane Katrina
Tsunami: JapanImpacts of climate changes
http://en wikipedia org/wiki/An Inconvenient Truth
1. 2 Importance
123752: Suchila, KKU, Thailand
200620052004 2011-4
Tsunami: Thailand Cyclone Nargis
Flooding: Thailand
https://www.google.co.th/search
http://en.wikipedia.org/wiki/An_Inconvenient_Truth
Earthquake Thailand (2014)
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New lifestyles and new technologies to avert more global warming: what’s the hurry?
It’s getting hotter
1. 2 Importance
123752: Suchila, KKU, Thailand
June, 2003: 15,000 excess deathsin France alone from heat wave
Source: V. Gutschick, NMSU (Emeritus)11
Rainfall distribution is changing…and becoming more extreme
…which may lead to more frequent droughts, and megadroughts, and wildfires
Source: V. Gutschick, NMSU (Emeritus)
1. 2 Importance
123752: Suchila, KKU, Thailand
Source: V. Gutschick, NMSU (Emeritus) 12
1
Why hurry?Sea level rise…in the long term
What’s needed already…will have to be multiplied manyfold
Ice caps are melting faster than originally predicted, from “unplugging”
1.2 Importance
123752: Suchila, KKU, ThailandSource: V. Gutschick, NMSU (Emeritus)13
Increased demand 50% by 2030 (IEA)
Energy
Climate Climate ChangeChange
1. Increasing population
2. Increasing levels of urbanisation
The Perfect Storm?The Perfect Storm?1. 2 Importance
123752: Suchila, KKU, Thailand
Water Increased demand 30% by 2030
(IFPRI)
FoodIncreased demand 50% by 2030
(FAO)
ChangeChangeurbanisation
3. The rightful goal to alleviate poverty
4. Climate Change
Source :John Beddington. 2009. Science in Government: Challenges for the Science in Government: Challenges for the 2121stst Century. Century. Campaign for Science and Engineering, London 10 December 2009.14
Recent effects of global warmingon agriculture and horticultural crops in Japan
• survey on the effect of global warming on agricultural industry in Japan
1.2 Importance
123752: Suchila, KKU, Thailand
http://www.agnet.org/library.php?func=view&style=type&id=20120104150721 15
White immature rice Cracked rice
1. 2 Importance
123752: Suchila, KKU, ThailandHinohikari Nikomaru (breeded in 2005)
http://www.agnet.org/library.php?func=view&style=type&id=2012010415072116
Rind puffing of Satsuma mandarin
Ch i th di t ib ti
1. 2 Importance
123752: Suchila, KKU, Thailand
Sunburned fruit Change in the distribution of suitable lands for apple
cultivation
http://www.agnet.org/library.php?func=view&style=type&id=20120104150721 17
1. 2 Importance
123752: Suchila, KKU, Thailand
Improvement of fruit quality by control of water status using plastic mulching and drip irrigation (fertigation) in citrus orchard
http://www.agnet.org/library.php?func=view&style=type&id=20120104150721
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1
Reducing water use,increasing yields
drip irrigation From 39% in 2009 to 52% in 2011• Australia, tomatoes using up to 70% less water
• USA tomato yield
1. 2 Importance
Resistant/tolerant varieties are
123752: Suchila, KKU, Thailand
• USA , tomato yield increase of over 25%.
• India, trials on gherkins save water 40%
• fungicide application rates reduced to 50%.
Source: Kamol Lertrat (2012)
necessary.
19
1.3 Principles and methods of breeding for resistance
Method for controlling disease and insect pests
o Quarantine
o Cultural controlไรตัวห้ํา
Source: รศ.ดร. นุชรีย์ ศิริ
1.3 Principles and methods
123752: Suchila, KKU, Thailand
o Chemical control
-Hyperparasites and predators
-Resistant varieties
o Integrated control
แมลงช้างปีกใส
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Host‐parasite Co‐evolution: is defined as the reciprocal adaptive
genetic change of two antagonists through reciprocal selective pressures.
Selection dynamics:
1.3 Principles and methods
123752: Suchila, KKU, Thailand
o Negative frequency dependent selectiono Over-dominant selection: Rr > rr, RRo Directional selection
http://en.wikipedia.org/wiki/Host%E2%80%93parasite_coevolution
21
Priorities in breeding objective for resistance to disease/insect:o Great amount of damage to the cropo Controlled at low cost by other means, does
Challenge facing the plant breeder
1.3 Principles and methods
123752: Suchila, KKU, Thailand
not merito Screening techniques is available (plant Patho/Ento) + Breeder
o Source of resistance (plant Patho/Ento) + Breeder
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oTo find out source of resistance:local adapted lines, breeder stock, exotic cultivars, wild and related species and created variation
Challenge facing the plant breeder (cont.)
1.3 Principles and methods
123752: Suchila, KKU, Thailand
created variationoMethods to introduce resistance into the existing breeding materialso Agronomic/quality accepted resistant variety, and do not neglect resistance to other pests.
23
2.1) Gene action and gene interaction
2.1.3 Genetics of host‐pathogen reactions
2.1.1 Concept of pathogen and host
2.1.2 Mechanisms of defense in plants against pests
Chapter II: Inheritance of resistance/tolerance
123752: Suchila, KKU, Thailand
2.2.1 World genetic resources
. .3 Ge et cs o ost pat oge eact o s
2.2) Sources of Resistance
2.2.2 Genetics resource management 24
1
Part II: Inheritance of resistance/tolerance (cont.)
2.3.1 Resistance breeding strategies
2.3) Heritability and breeding method
2.3.2 Breeding for specific resistance
123752: Suchila, KKU, Thailand
2.4.2 to environmental stress disease
2.4) Assessment of resistance/tolerance
2.4.1 to plant pests (bacteria, fungi and virus)
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Pathogen Host
Pathogenicity:
Avirulence vs virulence
o Pre-existing defense Mechanisms
2.1) Gene action and gene interaction:
2.1.1 Concept of pathogen and host
123752: Suchila, KKU, Thailand
Avirulence vs virulence
Non‐aggressive vs aggressive
Races vs pathotypes
o Infection induces defense mechanisms
https://www.google.co.th/search?q=potato+late+blight+pictures
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Pre‐existing defense
Mechanisms
o Cuticles
o Stomata
Pl t h i
Infection induces
defense mechanisms
o Periderm: plant cell accumulate lignin, turn to necrosis and inhibit disease growth
Mechanism of Resistance
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
o Plant hairs
o Phyto‐chemical:
‐tomatine anti Fusarium oxysporum
‐unsaturated lactones (glucosides in tulip anti F. oxysporum
o Formation of plant structure: gels and tylose inhibit disease epidemic
o Cell wall modifications: to protect plant cell from disease infection
o Hypersensitivities reaction
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• Immunity
• Disease escape
2.1.2 Mechanisms of defense in plants against pests
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
• Defense
• Resistance• Tolerance
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Resistance: no establishment of
Avoidance or escape: a mechanism that reduces the probability of contact between pathogen or insect pests and the plant
2.1.2 Mechanisms of defense in plants against pests (cont.)
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
Tolerance: the host develops, continues to grow, and produces well despite the pathogen’s presence
Resistance: no establishment ofthe pathogen in/on the host, or a limited establishment
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Proposed by H.H. Flor (1950)-study on flax rust “Gene for Gene Hypothesis” Virulence of a pathogen and the resistance of the host have a genetic
2.1.3 Genetic of host-pathogen reactions
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
resistance of the host have a genetic basis. For each gene that confers resistance in the host, there is a corresponding gene in the pathogen.
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1
Genetic Variation in Plant
1) Environmental Variation
2) Heredity variation:
Gene Action and Interaction
1) Gene action:
‐ Dominance
‐ Recessive
2) Gene Interaction:
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
‐ Quantitative Variation
‐Qualitative Variation
2) Gene Interaction:
‐ Complementary action
‐Modifying action
‐ Inhibiting action
‐Masking action (epistasis)
‐ Duplicate action
‐ Additive action31
1) Fungi:1.1 Obligate parasite
1.2 Facultative parasite
Variations: Mutation, genetic recombination
3) Bacteria:3.1 Gall forming bac.
3.2 soft rot bac.
3.3 Physiological or
Genetic Variation in Pathogen
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
genetic recombinationy g
metabolism altering bac.
Variations: binary fission, genetic recombination, mutation
2) Virus:
Variations: Mutation, genome recombination
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Type and Mechanism of Resistance
1) Base on heredity of resistance
1.1 Monogenic R.
1.2 Oligogenic R.
1 3 Polygenic R
3) Base on mechanism of R.
3.1 Active R. (Responsive R.)
3.2 Passive R.
4) Base on development stage of plant resistance
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
1.3 Polygenic R.
2) Base on epidemic of disease (Van de Plank)
2.1 Vertical R.
2.2 Horizontal R.
5) Base on population distribution
5.1 Quantitative R.
5.2 Qualitative R.
p
4.1 Seedling R.
4.2 Adult or mature R.
33
Types of Resistance
o Vertical resistance
o Horizontal resistance
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
http://en.wikipedia.org/wiki/Heritability
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• Hypersensitivity
• Specific resistance
• Major gene or oligogenic
Vertical Resistance
Resistance
Susceptibility
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
oligogenic
• Non‐durable or Qualitative resistance
1 2 3 4 5 6Races
Vertical Resistance to Races 2, 5, and 6
www.uky.edu/...breeding/.../Plant%20Breeding%20S07/bre...
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• Partial resistance
• Non‐specific resistance
Mi i t
Horizontal Resistance
Resistance
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
• Minor gene resistance
• Polygenic inheritance
• Quantitative resistance
Horizontal Resistance to all Races
1 2 3 4 5 6Races
www.uky.edu/...breeding/.../Plant%20Breeding%20S07/bre...
36
1
Genes in pathogen
R (resistant, dominant)
r (susceptible, recessive)
A dominant AR (‐) Ar (+)
Gene‐for‐gene reactions
Virulence(a) or Avirulence
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
a recessive aR (+) Ar (+)
(A) genes In thepathogen
37
Locks and Keys concept(Simplified by J.A. Browning)
Pathogen Host
R1R2 R1r2 r1R2 r1r2
A1A2 - - - +
For resistance:Host has a set of locks (R1R2) as dominant alleles
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
A1a2 - - + +
a1A2 - + - +
a1a2 + + + +
To establish disease: Pathogen has a set of keys as a virulence allele (a1, a2) to open the locks,or host much have no lock (r1, r2) for that keys
38
Genetics of Host Plant RelationshipGene for gene concept (Flor, 1956)
Genotype
Variety Plant Disease Symptom
Redwood NNPP AnAnApAp R
Genotype of flax and rust
2.1 Gene action and gene interaction
123752: Suchila, KKU, Thailand
Redwood NNPP ananApAp R
Polk NNpp AnAnapap R
Polk NNpp ananApAp S
Winowa nnpp All genot. S
Plant genot.: NNPP=R, Disease genot.: a=virulence; A=avirulence39
• Over 1750 genebanks exist world wide.
• About 7.4 million germplasm accessions
conserved in ex situ collections,
• More than 70% of the genetic diversity of
some 200-300 crops is already
2.2 Source of Resistance
2.2.1 World Genetic Resources
123752: Suchila, KKU, Thailand
Establishment of the Svalbard Global Seed
Vault, Norway a last resort safety back-
up repository of genetic resource
safeguard humanity.
conserved in genebanks (SBSTTA, 2010)
• Source: WIEWS 2009; Country
Reports; USDA-GRIN 2009, SOW2
40
USDA, TGRC
The Centre for Genetic Resources,
the Netherlands (CGN)
Active Genetic Resources for Pepper and Tomato
2.2 Source of resistance
123752: Suchila, KKU, Thailand
41
S. chilenseS. pennellii S. peruvianum
Genetic Resources for Tomato Improvement 2.2 Source of resistance
123752: Suchila, KKU, Thailand
© TGRCS. habrochaites
S. pimpinellifoliumS. l. var. cerasiforme
S. lycopersicum S. chmielewskii
Source: AVRDC 42
1
Resistance genes against insectsHessian Fly resistance genes H9, H13
Russian Wheat Aphid resistance genes Dn2, Dn4
• Most cultivars from the Pacific North-west carry only resistance gene H3
• RAPDs markers for H9: CCCAGTCACT & H13: TTGCTGGGCG
GrainGenes
(Source: Dweikat et al. 1997 TAG 94: 419-423)
D 2 7DS li k d t i t llit X 111 (3 M)
Grain Genes
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Green bug resistance gene Gb5
7A.7S-L7/5
Xpsr129
EcoR I
77SS
77AA
• Gb5: Large interstitial translocation from T. speltoides chromosome 7S-Linto chromosome 7A.
(Source: J. Dubcovsky et al. Crop Sci. 38: 1655-1660)
• Dn4:1DS between RFLP markers Xabc156 (11 cM) and Xksud14 (Source: Me et al 1998 genome 41: 303-306)Gran Genes
GrainGenes
• Dn2:7DS linked to microsatellite Xgwm111 (3 cM)
(Source: Liu et al. 2001 TAG 102: 504-510)
43
TVRC, KU:
2.2.2 Examples of genetic resources in Thailand
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Difficult to accessPBRCSA, KKU
Pepper 800 acc.
Tomato 1000 acc.
TVRC, KU:
Pepper 2827
acc. Tomato
acc.?
44
Active genetic resources for hot pepper and tomato
2000
8163
http://www.ars‐grin.gov/npgs/orders.html
Pepper Tomato websiteResources
www avrdc org
2.2 Source of resistance
123752: Suchila, KKU, Thailand
1011 1309 http://www.cgn.wageningen‐ur.nl/pgr
Banco de Germoplasma de
Hortalias (BGH; Brazil)www.bgh.ufv.br/
‐ http://tgrc.ucdavis.edu
www.avrdc.org
45
Germplasm*
elite line
Base/improve
Public/ Private
(Seed Co.)/ Grower
Food/health
food
Genetic
resources:
AVRDC
USDA
TGRC TVRC KKU
Seed co.
Col
lect
ion
Evaluation
Gene pool
Germplasm management at KKU
2.2 Source of resistance
123752: Suchila, KKU, Thailand
population
Pure/inbred lines
F1-hybrids
Pharmaceutical
products
(Animal) feed
additive
etc.
Selection
Selection
Svalbard Global Seed Vault
46
Collection
Evaluation
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Evaluation
Distribution
47
Examples of resistant and other characteristics sources of pepper and tomato
Phyto-nutrient Capsiate, pigment Lycopene,
ß-carotene
Disease Anthracnose Bacterial wilt
Trait Pepper Tomato
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Disease
resistance***
Anthracnose,
Gemini virus,
Begomo virus
Bacterial wilt,
TYLCV,
Begomo virus
Others
(Agronomic traits)
Male sterile Raisin tomato
48
1
Evaluation and characterization
Evaluation
100 acc./year
KKU accession number KKU-P 11008Duplicate number -Temporary number -Variant -Species Capsicum annuum L.Subtaxa -Pedigree / Cultivar Name Jinda-YaoCountry Thailand/KKU
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Location acc./year
KKU 40
U. Ratchapat CM. 30
U. Ratchapat MSK. 30
(total 300 acc.)
CountryCharacterized year and Rainy Season, 2006Remarks -
Desctiptor Name Value Desctiptor Name Value
Cotyledonous leaf shape - Stem color Green
Cotyledonous leaf color Green Root distribution Low
Hypocotyl pubescence -
Lyfe cycle Biennial Leaf colour Green
Stem color Green with purple stripes Leaf shape LanceolatePlant height (cm.) - Mature leaf length (cm.) 8.5Plant canopy width (cm.) - Mature leaf width (cm.) 3Plant growth habit Intermediate
Days to flowering 40 Anther colour PurpleNumber of flowers per axil One Male sterility AbsentFlower position Erect Calyx annular constriction AbsentCorolla colour White
Fruit colour at intermediate stage Green Neck at base of fruit AbsentFruit colour at mature stage Red Fruit shape at blossom end PointedFruit shape Elongate Fruit surface SmoothFruit length (cm.) 7.5 Fruit pungency High (~ 152,995.80)Fruit width (cm.) 1.4 No. of locules -Fruit shape at pedicel attachment Obtuse
Seed colour Straw 100-seed weight [g] 0.502Seed surface Smooth Number of seeds per fruit -
No. of fruit 300 Dry weight (g) -Fresh weight (g) 530 Remark -
Last update : March' 2007
Seedling Data
Vegetative Data
Inflorescence Data
Fruit Data
Seed Data
Yield Data
49
Germplasm distribution
Field day/Varietal trials
at KKU
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Hot pepper in rainy season Tomato in dry season
50
Stock Maintenance:
Cold room conditions
• Medium-term conditions : under refrigeration at 5-10°C.
• All seed samples sealed in a suitable air-tight container in
%
•Drying at 10-25 ° C and 10-15 %RH
• Drying at 5-20 ° C and 15-25% RH
2.2 Source of resistance
123752: Suchila, KKU, Thailand
Viability monitoringstorage environment RH 15±3%.
•Germination should exceed 85% & 75 % some
vegetables
•within 12 months after receipt of the sample Regeneration
•Viability drops below 85 % of the
initial viability.
51
Data Base ManagementDatabase QueriesAccessions... Geographic Data on
Wild Species Accessions... Core collectionGenesList of Gene Names and Symbols...View Naming Rules
Example of seed request
2.2 Source of resistance
123752: Suchila, KKU, Thailand
gImages ColleaguesOther ResourcesRecent AcquisitionsTop 20 most requested Accessions (2011). Recent database/website updates:(Google maps now available for wild species)(All Wild species localities available)
Solanum lycopersicoides introgression linesSolanum habrochaites (L. hirsutum) introgression lines2nd generation pennellii ILs2005 collections from Chile2001 collections from ChileSeed Request InformationHow to Request SeedContact Information 52
Heritability :is a concept that summarizes how much of the variation in a trait is due to variation in genetic factors. This term is used in reference to the resemblance between parents and their offspring. In this context, high heritability implies a strong resemblance between parents and offspring with regard to a specific trait while low heritability implies a low level
2.3 Heritability and Breeding Methods
123752: Suchila, KKU, Thailand
regard to a specific trait, while low heritability implies a low level of resemblance.
Figure 1: Heritability estimation.Low (panel a) and high (panel b) heritability can be estimated from the regression (h2) of offspring phenotypic values vs. the average of parental phenotypic values.© 2008 Nature Education All rights reserved
53
Heritability Broad-sense heritability, defined as h2
b = VG/(VG +VE)
Narrow-sense heritability, defined as h2 = VA/(VA + VD + VE)
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
defined as h n VA/(VA VD VE)
Figure 3. Strength of selection (S) and response to selection (R) in an artificial selection experiment,h2=R/S
Figure 4. Sir Francis Galton's (1889) data showing the relationship between offspring height (928 individuals) as a function of mean parent height (205 sets of parents).
http://en.wikipedia.org/wiki/Heritability54
1
Specificity in the parasite:
2.3.1 Resistance Breeding Strategies
Physiological race: Pathogen genotypes share a group of cultivars
Differential hosts (cultivars): Using differential cultivars toidentify physiological races
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
share a group of cultivars to which they are virulent
identify physiological races or vice versa.
Cultivars Races
R1 R2 R3 R4
C1 - + - +
C2 - - + +55
Tomato Late Blight Differentials in AVRDC Taiwan
Host Differentials
Pathogen race Ph+,
Ph-1
WV700
(Ph-2)
CLN2037
(Ph-3)
L3708
(Ph-3 + Ph-4)
LA1033
T1 S R R R R
T1,2 S S R R R
T1,3 S R S R R
T1 2 3 S S S R R
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
T1,2,3 S S S R R
T1,3,5 S R S R S
T1,2,3,4 S S S S R
T1,2,3,5 S S S R S
T1,3,4,5 S R S S S
T1,2,3,4,5 S S S S S
Chen et al., 2008 R= host gene confers resistance to that pathogen race S= susceptible reaction. Pathogen race overcomes resistance
56
Individual Major Genes
2.3.2 Breeding Methods for Specific Resistance
Application of gene pyramiding
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
Using multi-lines
57
Individual Major Genes
Breed cultivars with major genes that control the
l t t
2.3.2 Breeding Methods for Specific Resistance (cont.)
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
prevalent pest races
Select progeny from a segregating population or
Transfer major genes from other sources
58
Application of gene pyramiding
Put all knownmajor genes into one line
Analysis of genome
2.3.2 Breeding Methods for Specific Resistance (cont.)
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
Singh A et al. AoB PLANTS 2012;2012:pls029
Published by Oxford University Press on behalf of the Annals of Botany Company.
Analysis of genomeintrogression associated with resistance genes/QTL. (A) xa13 on chromosome 8 and Xa21, Pi54and qSBR11‐1 on chromosome 11 (B) in ‘Pusa1608’ families.
59
Ty-1
Ty-3
Chromosome 6 Chromosome 11 Chromosome 3
Mapped TY Resistance Genes in Tomato at AVRDC
Ty-5
Chromosome 4
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
Ty-2
Ty-4
Source: Peter Hanson, AVRDC 60
1
Using multi-lines
Develop many individual lines each with individual major resistance genes (isolines (or
2.3.2 Breeding methods for Specific Resistance (cont.)
2.3 Heritability and breeding methods
123752: Suchila, KKU, Thailand
near isogenic lines))
Mix the seed of these lines together to get multilines
Protection against a broad spectrum of races
61
2.4) Assessment of Resistance and Tolerance
2.4.1 Concept of evaluation for resistance/tolerance
2.4.2 Assessment for plant pests (bacteria, fungi and virus) resistance
123752: Suchila, KKU, Thailand
2.4.3 Assessment for environmental tolerance
fungi and virus) resistance
62
under:
o Field condition
o Hot spot condition
A tifi i l/l b t
Resistance levels of disease assessment
o Disease incidence
o Disease severity
2.4.1 Concept of evaluation for resistance or tolerance
Selection/assessment for resistance
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
o Artificial/laboratory condition
o Disease severity
63
Gene controlling for agronomic traitsand resistance to biotic and abiotic stresses are expected to be located at different regions (Blum, 1980)…
2.4.1 Concept of evaluation for resistance or tolerance
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Therefore, assessment the responses of different varieties should be done under separate and suitable conditions, and particular generations.
64
Screening diagram Assessment for resistant to diseases-insect pests and environmental stresses (Adapted from Blum, 1980)
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
When?: Generations, seasons
Where?: Environmental locations or conditions
How? Inoculations and evaluations
65
2.4.2 Assessment for plant pests resistance
Assessment for bacteria responses
Assessment for fungi responses
Assessment for virus responses
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
66
1
Breeding tomato varieties for bacterial wilt resistance
Assessment for bacteria responses
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
67
Bacterial wilt
Ralstonia solanacearum
Serious disease and major constraint in
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
j
the tomato production
Reduce field stand 30-100 %
68
Tomato Bacterial wilt
Properties
-Soil borne disease
-wide host range
-No effect to pesticide
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Characteristics of bacterial wilt resistance
-Location specific
-Strain/temperature dependent
-Not immune: internal bacterial population correlated to
symptom
Wang et al, 2000a; Wang et al, 2008
69
The International set of resistance
source in tomato to bacterial wilt
worldwide evaluated 33 var.
with 1 susceptible check
3 sources of resistance;
Hawaii, Philippine and North Carolina? (Scott et al., 2005)
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Hawaiian resistance source:
•a broad-based and the most stable and durable
resistance
•HW7996 the best source of partial resistance to race 3
Wang et al., 1998; Carmeille et al., 2006
70
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
71
Symptom of wilting caused by
Ralstonia solanacearum
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand72
1
Some screening techniques at KKU
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
73
Materials and methods
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
74
Table 1. β-carotene content,
fruit yield and some
characteristics of 34 tomato
cultivars grown under field
and plastic-net house
conditions during October
Thirty tomato cultivars from many resources, two resistant and two susceptible
were screened for bacterial wilt resistance.
123752: Suchila, KKU, Thailand
2008-Febuary 2009 at the
experiment field, Khon Kaen
University, Thailand.
75
Table 2. Responses of fruit yield and bacterial wilt responses in 32 tomato lines/cultivars under the concrete block
with bacterial wilt inoculums during dry season (October 2008-Febuary 2009) and responses to bacterial wilt
disease of 12 tomato cultivars under 2 seasons, dry (October 2008-Febuary 2009) and rainy season (May-
September, 2009) at the experimental field, Khon Kaen University,Thailand.
Bacterial wilt responses and fruit yield were evaluated.
123752: Suchila, KKU, Thailand
76
Preparation of bacterium inoculums
R. Solanacearum : 3 Isolates ()
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Inoculum concentration = 0.2
(107-10
8cfu/g. soil mixed population)
77
Seedling preparation
Seedling: 25-30 days
after sowing
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
RCBD 3 reps./variety
78
1
Inoculation Methods
1. Root inoculation
2. Micropipette (micropipette technique)
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
3. Scalpel leaf clip method or leaf clipping
4. Soil drenching method
79
Scalpel leaf clip method or leaf clipping
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Micropipette Soil drenching method80
Pathogen morphology and Type of Symptoms
OOZES COLONY
123752: Suchila, KKU, Thailand
STEM CROSS SECTION
81
Evaluation
Rating score
• 1 : No symptoms
• 2 : One leaf wilted
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
• 3 : Two to three leaves wilted
• 4 : Four or more leaves wilted
• 5 : Whole wilted (dead plant)82
Wilting intensity computing
% Wilt intensity : %I
I = [ ∑ (Ni x Vi) / (N x V )]x 100
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
∑Where Ni mean : number of plant wilt respective disease rating
Vi : disease rating (1,2,3,4 or 5)
V : the highest disease rating
N : the number of plant observed
Winstead and Kelman (1952)
Chanh (1989)
83
RM
R
M
SS
20 40 60 80
H
S
100
Highly
resistant
123752: Suchila, KKU, Thailand
R S S
Bacterial wilt phenotype
84
1
Varietie
s
% dead plant % IntensityWilt intensity observation
(Phenotype)
Week 1 Week 2 Week 3 Week 4 Week 1 Week 2 Week 3 Week 4 Week 1 Week 2 Week 3 Week 4
V1
V2
V3
V4
V5
Data recorded
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
V5
V6
V7
V8
V9
V10
85
Genotypic analysis
Materials and methods
Dellaportaextraction method
Dellaportaextraction method
123752: Suchila, KKU, Thailand
200 plants of F2 200 plants of F2
6% poly-acrylamide gel6% poly-acrylamide gelVisualized with the gel
document photographerVisualized with the gel
document photographer
M P1 P2 Homozygouse S
Homozygouse R
Heterozygous
Homozygouse SHomozygouse RHeterozygous
F2
86
DNA Marker linked to Bacterial wilt resistance on chromosome 6 and 12
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
500 bp
87
M P1 P2
Homozygouse SHeterozygous
F2
Statistical analysis
Segregation ratio by χ2 test =(O‐E)2/E
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Homozygouse R
Homozygouse S
Homozygouse R Heterozygous
88
Data recorded
1 1 R1 Hawaii7996 RR RR2 2 L1 PI114968 84 AI SS SS3 3 F1 L X R1 RR RR4 4 F2 L X R1 SS5 5 F3 L X R1 RS RS6 6 F4 L X R1 SS RR7 7 F5 L X R1 RR RS8 8 F6 L X R1 SS SS9 9 F7 L X R1 RS RS
10 10 F8 L X R1 RS SS
No.DNA
No.CCA code
PedigRee name SLM6-17 SLM12-2
Data were analyzed by chi-square
test to as certain the goodness-of-
fit between the expected ratio
following by Mendelian ratio
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
10 10 F8 L X R1 RS SS11 11 F9 L X R1 RS RR12 12 F10 L X R1 RS RS
Genotype Expected frequency No. of plant Number of plant
χ² P Resistant Susceptible
Hawaii 7996 F2 3:1 200 F2 15:1 200 F2 13:3 200
following by Mendelian ratio
89
Breeding hot pepper for anthracnose resistance
Assessment for fungi responses
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
90
1
Host : Susceptible cultivars ( Variety and Fruit Stage )
Factors affecting anthracnose disease
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Pathogen
:
91
Year Author
species of
capsicum spp.
Resistant source Country reported
Cc Cg Ca
1999 AVRDC C. baccatum PBC81, PBC80 Taiwan
C. chinense PBC932
2003 Yoon C. baccatum PBC81, PBC80 Korae
2004 Yoon etal., 2004 C. baccatum PBC81, PBC80 Korae
2004 Voorrips et al. C. chinense PBC932 Korae
2007 Kim et al. C. baccatum PBC81, PBC80 Korae
C. annuum Daepong-cho
Source of anthracnose resistance
123752: Suchila, KKU, Thailand
2005 Pakdeevaraporn et al. C. chinense PBC932Thai land
C. baccatum PBC81, PBC80
2005 Park C. baccatum PI594137 Korea
2006 Lin et al. C. chinense PBC932
2008 Prasath C. baccatum PBC81
2005 Pae et al C. baccatum PBC1430
PBC1439
PBC1478
PBC880
C. annuumTCO6903 (R95)
92
Some resistant sources in hot pepper varieties
Progressive lines derived from PBC 932 and PBC 80
123752: Suchila, KKU, Thailand
Original resistance sources
93
FactorsFruit development
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
94
C gloesporioides
Pathogen
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
C. capsiciC. capsiciC. acutatum
C. gloesporioides
Than., 2008
Different species of Colletotrichum grown on PDA
Cc930 Cc1141 Cc388 Ca524 Ca153
95
Pathogen
Cc‐cap26 Ca_MUJ5
- 500 spore/ml
-2 ul, RH 98%, Dark
Microinjection Method
Inoculums preparation for inoculation and evaluation
123752: Suchila, KKU, Thailand
- Evaluation 5 DAI
Following Park et al (1999) technique96
1
Green mature and ripe mature fruit stage for inoculation
123752: Suchila, KKU, Thailand
Mature fruit: about 35 and 45 day after anthesis
97
Evaluation of disease reactions
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
98
score Resistant level Symptom details
0 HR, highly resistant No infection
1 R, resistant 1-2% of fruit area shows necrotic lesion or a larger water-soaked lesion surrounding the infection site
3 MR d l i 2% f h f i h i l i li b
1) Percentage of necrotic lesion on fruit: six rating scores
Two techniques of disease evaluations
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
3 MR, moderately resistant >2% of the fruit area shows necrotic lesion, acervuli may be present, or water-soaked lesion up to 5% of the fruit surface
5 MS, moderately susceptible
>5-15% of the fruit area shows necrotic lesion , acervuli present , or water-soaked lesion up to 25% of the fruit surface
7 S, susceptible >15-25% of the fruit area shows necrotic lesion with acervuli
9 HS, highly susceptible >25% of the fruit area shows necrosis, lesion often encircling the fruit, abundant acervuli
Montri et al., 2009 99
2) Dimension of necrotic lesion on fruit (1 mm ‐36 mm): two rating scores R S
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
1) Resistance: necrotic lesion < 4 mm.
2) Susceptible: necrotic lesion > 4 mm.
AVRDC
100
No. Rep box variety lesion green red
1 1 1 1
2 1 1 2
3 1 1 3
4 1 1 4
5 1 1 5
6 1 1 6
7 2 1 1
8 2 1 2
Data collections
Data sheet
2.4 Assessment of resistance and olerance
123752: Suchila, KKU, Thailand
8 2 1 2
9 2 1 3
10 2 1 4
11 2 1 5
12 2 1 6
13 3 1 1
14 3 1 2
15 3 1 3
16 3 1 4
17 3 1 5
18 3 1 6
Statistic analysis by RCBD, Test significant at p=value 0.05
and 0.01
101
Data analysis
Disease
reaction1/
Disease
reaction1/
101
102
103
104
105
variety
C. acutatum_MUJ5
Ca - green mature Ca - ripe mature
disease incidence disease incidence
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
106
107
108
109
110
111
susceptible checkResistant check Conclusion for Disease reactions
102
1
Molecular marker aids selection
Distribution of marker
for anthracnose resistance
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
103
DNA extraction and gel electrophoresis
2.4 Assessment of Resistance and Tolerance
123752: Suchila, KKU, Thailand
1) Harvest the younger leaves
2) Extraction DNA
3) PCR and running gel (Acrylamide gel)
4) Scoring DNA banding104
genotype Primer name Sequencings Linked
Forward primer Reverse primerC.capsici HpmsE143 CCATTCAGCTAGGGTTCAGTCCA CGACCAAATCGAATCTTCGTGA 9
Anth9-CAPS Unpublished Unpublished 9
C.acutatum HpmsE032 ATGCGCAAAGGGAGAAAATTCA CGAACTAACCGTTCATGGTGGA 12
Primers
2.4 Assessment of Resistance and Tolerance
123752: Suchila, KKU, Thailand
Anth12-CAPS Unpublished Unpublished 12
SCAR-Indel 100 bp GGTATCTTATTTCATAGGGACCAGGCA TTTGCGGTAGTGACAACAACTTTACAGCCA -
105
Scoring of DNA banding pattern
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Lane 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
DNA score 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3
Lane 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43
DNA score 3 1 3 1 3 1 3 1 3 1 3 3 3 3 3 1 3 3
106
Evaluation of Yellow Tomato Leaf Cure Virus (TYLCV)
Assessment for virus responses
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Source: AVRDC, 2012
107
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
108
1
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
109
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Source: AVRDC, 2012 110
Mapped TY Resistance Genes in Tomato
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
111
Tomato BGVsThe World Vegetable CenterThe World Vegetable Center
AVRDCAVRDCHost range of TYLCV2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
112
Pepper BGVs
Host range of TYLCVThe World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
113
Pepper BGVs
Host range of TYLCV
The World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
GV
114
1
Cucurbit BGVs
SquashSquash
Host range of TYLCV
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, ThailandBottle gourdSquash plantsChayote The World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC 115
Cucurbit BGVs
Host range of TYLCV
The World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Ridge gourdCucumber 116
Legume BGVs
Host range of TYLCVThe World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Mungbean Yard long bean
Soybean
117
Okra BGVs
Host range of TYLCVThe World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
118
Eggplant BGVs
Host range of TYLCVThe World Vegetable CenterThe World Vegetable Center
AVRDCAVRDC
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand119
TYLCV disease vector and transmission
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
120
1
Genetic resource from AVRDC
35 accessions of pepper varieties
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
121
Materials and Methods
V1 V2 V3 V4 V5 V6 V7 V8
V9 V10 V11 V12 V13 V14 V15 V16
V17 V18 V19 V20 V21 V22 V23 V24
LP
LP
LP
LP
Experimental design with 56 tested varieties and local variety
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
V25 V26 V27 V28 V29 V30 V31 V32
V33 V34 V35 V36 V37 V38 V39 V40
V41 V42 V43 V44 V45 V46 V47 V48
V49 V50 V51 V52 V53 V54 V55 V56
LP
LP LP
LP
LP
LP
LP: Local check
V1-V56: Tested varieties
122
Material and methods: Field evaluation for TYLCV
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
123
Evaluations of TYLCV: plant symptoms
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
124
Evaluations of TYLCV: leave symptoms
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
125
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
126
1
Percentage of disease reactions
% Disease
symptoms= Σ
(disease score x plant no.)
Total plant no. x maximum disease score
x 100
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
127
LP 1 2 3 4 5 6 7 8 9 10 11 12 LP LP number 1 2 3 4 5 6 7 8 9 10 11 12 LP
1 105 121
2 127 117
1 126 122
2 133 129
1 114 116
2 128 103
1 124 136
1
2
3
4
Symptom rating (visual) Replication: I Date (day/month):
Block sub block row number
Plant No.
Test plants Test plants
Table for data recorded
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
2 130 131
1 134 132
2 111 119
1 110 135
2 118 112
1 306 120
2 113 109
1 107 104
2 101 115
1 108 102
2 123 125
1
4
5
6
7
8
9
128
The Responses of Hot pepper to Drought Stress
2.4.3 Assessment for environmental tolerance
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
129
Chilii pepper production in Thailand
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
low yield and low qualitylow yield and low quality
80% under rain-fed area
130
400
500
600
700
800
900
Nakorn Rachasima
Chaiyaphum
Loei
Norteast of Thailand
ainfal
l (mm.
)
Fluctuation of rainfall (mm.) in chiili pepper commercial
production area of Northeast of Thailand
123752: Suchila, KKU, Thailand
0
100
200
300
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Ra
(a) (b) (c)
Note: (a) vegetative stage, (b) flowering stage and (C) fruiting stage 131
How to improve high pungency, high yield and
produced in short-term drought stress condition?
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Breeding Management
132
1
Relation of water in soil, plant and atmosphereAtmospheric water
Transpiration : stomata opening, RH
or PD
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Soil water
Water in Plant Absorption : root penitration, moisture
tension of soil
TRANSLOCATION
133
Soil moisture measurement• Gravitational method
soil moisture percentage = (wet soil weight - dry soil weight) x 100/ dry soil weight
• Tensiometric method
2.4 Assessment of resistance and tolerance
http://www.calafrica-sa.co.za/
123752: Suchila, KKU, Thailand
m m
http://www.calafrica-sa.co.za/http://www.calafrica-sa.co.za/soil_moisture_measurement.htm 134
• Gypsum block
Soil moisture measurement measurement2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
• Neutron probe Hydro probe
http://aces.nmsu.edu/pubs/_h/H640/welcome.html
http://www.agw.kit.edu/english/field_instruments.phphttp://www.pavetesting.com/hydroprobe.php
http://cals.arizona.edu/pubs/water/az1220/
135
Plant water status measurement
Relative water contentRWC = (Fresh weight - Dry weight)/
(Fully turgid weight - DW) X 100
Leaf water Potential
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Leaf water PotentialPressure chamber (Kg/cm2)
1 Ib/In2 = 6.895 x 103 Pa
1 Kg = 2.2046 Ib
1 Ib = ½.2046 Kg
1 In = 2.54 cm
½.2046
Kg/(2.542)
= 6.895 x 103 Pa
1 Kg/cm2 = 0.09807 MPa136
Water Potential in Plants
Determine water‐deficit stress in plants.
Determine drought tolerance in plants,
The irrigation needs of different crops
How the water status of a plant affects the quality and yield of plants
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
How the water status of a plant affects the quality and yield of plants.
Pure Water (no solutes) at normal pressure and elevation where, Ψw = 0
Ψw is increased by an increase in pressure potential (ΨP)
Ψw is decreased by addition of solutes which lowers the solute potential
(ΨS )
137
•Stress types: water (drought and flood), temperature (high temperature
and energy balance, chilling and freezing), irradiance, others
•Level of the stress:
Severe, moderate and mild
•Period of the stress:
Treatment application techniques
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Prolong (gradual), short term or transient (sudden)
•Application technigues:
On farm vs Lab. Research:
Drought: lab (water application vs PEG)
Temperature: shading vs Phytotron room or green house
Salinity: saline soil vs Lab
138
1
Experimental design- conducted under greenhouse condition- RCB design with 4 replications- varieties were divided into two groups;
1st gr. were treated with drought stress2nd gr. were control treatment (optimum water applied)
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand139
Drought treatment
1-10 DAF100%
11-20 DAF75%
21-30 DAF50%
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
75% 50%
31-40 DAF25%
-no watered-After severe wilting overnight (re-watered)140
Measurement of water status by Pressure Chamber
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
• Quick method, commonly used
• Equipment can be used in the field; can
be heavy and bulky
• Dangerous pressure levels are applied in
the chamber Source: Bowers and Sterling, Department of Entomology, Plant Pathology, and Weed Science, NMSU, 2004
141
The positive pressure reading from the plant tissue tested in the
previous slide was 7 bars, a stressed plant.
To estimate the water potential, we must first convert the positive
pressure from bars into MegaPascals (MPa).
Water potential measurement
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
10 Bars = 1 MPa
7 Bars = 0.7 MPaYP(air) + YP(xylem) = 0
pressure from bars into MegaPascals (MPa).
Ten bars is equal to one MegaPascal, so 7 bars equals 0.7
MegaPascals.
142
60.00
70.00
80.00
90.00
100.00
LWP
(M
Pa
)R
WC
(%
)
KKU-P11039 KKU-P13049 KKU-P11015
Control
Stress
Control
Stress
**
****
** *****
****
-1.00
-0.50
-
LWP
(M
Pa
)
Control
Stress
** ****
**- RWC and LWP of the treated plant were decreased at 21 30 d 31 40 DAF ti l ( d t th
Water status
-1.50
-1.00
-0.50
-
**
123752: Suchila, KKU, Thailand
-1.50
60.00
70.00
80.00
90.00
100.00
RW
C (
%)
Control
Stress
** ****
**
** **
** *
-1.50
-1.00
-0.50
-
60.00
70.00
80.00
90.00
100.00
LWP
(MP
a)
Control
Stress
Control
Stress
** **
*****
KKU-P12010 TakanotsumeKKU-P11031 KKU-P28006
KKU-P31118
KKU-P33030
21-30 and 31-40 DAF, respectively (compared to the control plant)
143
Plant response to drought stress
Physiological response• stomaltal closure reduce photosynthesis• Leaf development: small, mature and fall.• Produce hormone: ABA, Ethylene ,
Morphological response
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Morphological response• Increase leaf thickness and low leaf no. • Low shoot: root ratio• Chlorophyll damage• Leaf roller and wilt
144
1
Cultivars
Yield Growth
Fresh Weight (g) Dry Weight (g) Fruit Length (cm.) Fruit width (cm.) Plant Height (cm.) Stem diameter (cm.)
C S C S C S C S C S C SLow
KKU-P11039 639.96 368.24** 127.99 66.28** 3.74 3.73ns 5.62 5.49* 121.25 120.08ns 1.60 1.34**
KKU-P13049 241.04 178.27** 69.90 51.70** 0.38 0.37ns 3.22 3.18ns 151.85 136.78* 1.99 1.65**
KKU-P11015 503.12 268.00** 120.75 58.96** 1.54 1.52ns 7.30 7.09** 167.83 161.25ns 1.74 1.53**
Medium
Growth and yield responses
- Plant height and Stem diameter were decreased (5-20% and 10-30% of control) in the treated plant dependent on
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
KKU-P12010 385.94 317.05ns 119.64 98.28ns 0.51 0.50ns 3.62 3.56ns 128.88 138.55* 1.44 1.24**
KKU-P11031 400.17 325.71** 128.05 104.23** 0.58 0.57ns 4.32 4.18* 146.50 133.25* 1.66 1.66**
KKU-P28006 644.41 363.73** 161.10 87.29** 1.72 1.65ns 7.77 7.53** 155.45 139.50** 1.72 1.48**
Takanotsume 165.56 118.15** 41.39 28.36** 2.21 2.20ns 8.38 6.20** 125.31 103.78** 1.22 1.13ns
High
KKU-P31118 318.05 253.71ns 89.05 71.04ns 0.55 0.53ns 2.84 2.80* 144.84 140.25ns 1.44 1.29**
KKU-P33030 356.87 297.94** 96.35 80.44** 0.68 0.67ns 2.77 2.44** 120.55 111.9ns 1.68 1.13**
in the treated plant dependent on varieties- Fruit fresh weight and dry weight were decreased (20-50% of control) in the treated plant dependent on varieties
145
Water Use Efficiency ; WUE
Photosynthetic rate/transpiration of leaf
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Total dry mass/total transpiration
Total yield/(water use x HI)
146
EvapWUE
RWCWSDDWWSat
FWWSatWSD
DWWSat
DWFWRWC
mpw
2
2kgorginlossOH
gormginuptakeCONet
5YieldCroporMatterDry
.inUsedWater
4100
3100.
.
2100.
1
EQUATIONS AND ABBREVIATIONS
Abbreviation
w = Water Potential RWC = Relative Water Content ra = Atmospheric Resistance = Osmotic Potential WSD = Water Saturated Deficit rs = Stomatal Resistance p = Turgor Potential WUE = Water Used Efficiency rm = Mesophyll Resistance m = Matrix Potential D = Diffusivity PS = Photosynthesis Trans. = Transpiration Evap. = Evapotranspiration
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
rmrsra
rsra
OH
COWUE
DOH
Drmrsra
OHOHTrans
DCO
Drmrsra
COCOPS
OH
extint
OH
extint
2
2
2
22
2
22
64.0
76
7resistance
)][]([.
6resistance
64.0
)][]([
2
2
147
Example of drought stress experiment
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148
Fig. 1. (A–C) Monthly mean air temperature, relative humidity and light intensity under a plastic net house
and (D) monthly mean rainfall under field condition at the experimental farm at Khon Kaen University
during the rainy season (May to Oct. 2009).
E i t l diti d i t i d
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Environmental conditions during stress periods
149
Fig. 2. The values of leaf water potential response at 10-d intervals of the control (C ) and drought-
stressed plants (T ) in nine hot pepper cultivars.**Significant difference between control and drought
treatments within cultivar at P # 0.01 level by t test.
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Different species showed different water status under drought stress
150
1
Table 2. Leaf water potential (LWP), relative water content (RWC), leaf area (LA), and specific leaf area
(SLA) between the control plant (C) and drought-stressed plant (T) of nine hot pepper cultivars at 31 to
40 d after flowering.
Different species showed different
123752: Suchila, KKU, Thailand
Table 3. Plant height, stem diameter,
and shoot-to-root ratio between the
control plants (C) and drought
stressed plants (T) of nine hot pepper
cultivars at second harvest.
pgrowth development under drought stress, compared to their control treatment
151
Table 4. Yield and fruit quality between the control plants (C) and drought-stressed plants (T) of nine
hot pepper cultivars.
Table 5 Capsaicin (Cap) dihydrocapsaicin (Di) capsaicinoid (CAPS) and capsaicinoid yield (CAPS yield) Different species showed different
123752: Suchila, KKU, Thailand
Table 5. Capsaicin (Cap), dihydrocapsaicin (Di), capsaicinoid (CAPS), and capsaicinoid yield (CAPS yield)
between the control plants (C) and drought-stressed plants (T) of nine hot pepper cultivars.
pYield and pungency under drought stress, compared to their control treatment
152
Cultivar PAL (μmole mg-1
protein) C4H (AU min-1
g-1
) CS (μmole mg-1
protein) POD (AU min-1
g-1
)
C1/
D1/,2/
% increase3/
C1/
D1/,2/
% increase3/
C1/
D1/,2/
% increase3/
C1/
D1/,2/
% increase3/
Yuyi 33.99d 74.02e** 217.8 1.092b 1.167c* 106.9 67.6f 119.6g* 176.9 16.50a 28.13ab* 170.5
Huay-Siiton 92.59bc 135.59b* 146.4 1.181a 1.298a* 110.0 104.9ef 316.7de** 301.9 14.25ab 22.26bc* 156.2
K
Table 2. Phenylalanine ammonialyase (PAL), cinnamic-4-hydroxylase (C4H), capsaicin synthase (CS) and peroxidase (POD)
activities under a control (C) and a drought stress (D) treatments
Bio-molecular Traits for Drought Stress
Different species showed different
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
Keenoo-
Pama
67.68c 109.16bc* 161.3 1.084b 1.215b* 112.2 546.8c 628.5bc 114.9 1.26f 2.46g* 195.2
BGH 1719 148.91b 165.23ab 110.9 1.177a 1.214b* 103.2 752.4ab 578.3c 79.9 5.06d 31.31a** 618.8
Perennial 173.54a 188.32a 108.5 1.093b 1.191b* 109.0 936.4a 982.7a 104.9 3.13de 20.63c** 659.1
Mean 97.34 134.46 154.6 1.125 1.217 108.2 481.6 525.2 155.1 8.04 20.96 360.0
enzymes activities under drought stress, compared to their control treatment
153
Bio-molecular Traits Under Drought Stress
Figure 1. Sum of Cap and Di (CAPs) which
were redrawn from Phimchan et al9
and
activities of phenylalanine ammonia-lyase
(PAL), cinnamic-4-hydroxylase (C4H),
capsaicin synthase (CS) and peroxidase
(POD) in hot pepper fruit between non-
drought and drought stressed plants.
L 1 M 1 M 2 H 1 H 20
20000
40000
60000
80000
100000
120000
140000
160000
180000
Non-drought stressed plantDrought stressed plant
Capsaicinoids (CAPs)
CA
Ps
(SH
U)
Cultivars
150
200
250
Non-drought stressed plantDrought stressed plant
Phenylalanine ammonia-lyase (PAL)
e m
g-1
prot
ein)
Cinnamic-4-hydroxylase (C4H)
U m
in-1
g-1 )
1.0
1.5
2.0
2.4 Assessment of resistance and tolerance
123752: Suchila, KKU, Thailand
L 1 M 1 M 2 H 1 H 20
50
100
PA
L (u
mol
e
Cultivars Cultivars
L 1 M 1 M 2 H 1 H 2
C4H
(A
U
0.0
.5
Capsaicin synthase (CS)
L 1 M 1 M 2 H 1 H 20
200
400
600
800
1000
1200
CS
(um
ole
mg-
1 pr
otei
n)
CultivarsL 1 M 1 M 2 H 1 H 2
0
10
20
30
40
Peroxidase (POD)
PO
D (
AU
min
-1g-
1 )
Cultivars
Only phenylalanine ammonia-lyase (PAL) showed positive association
to capsaicinoids contents under drought stress
Source: Pimchan et al, 2014 154
Experiments: 3 chilli pepper cultivars, 3 replications
and 3 plants per replication 3 levels of drought stress treatment
Exercise for drought stress
2.4 Assessment of resistance and tolerance
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Data recorded: LWP, RWC, plant height, plantcanopy, Stem diameter and SPAD chlorophyllreading
155
Cultivar Rep.LWP RWC
T1 T2 T3 T1 T2 T3
Bhut jolokia
1
2
3
Table 1 Leaf water potential (LWP) and relative water content (RWC) of 3 hotpepper cultivars treated by 3 levels of drought stress
2.4 Assessment of resistance and tolerance
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Akanee pirot
1
2
3
Habanero
1
2
3
156
1
Cultivar Plant no.Plant height (cm.) Plant canopy (cm.)
T1 T2 T3 T1 T2 T3
Bhut jolokia
1
2
3
Table 2 Plant height and plant canopy of 3 hot pepper cultivars treated by3 levels of drought stress
2.4 Assessment of resistance and tolerance
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Akanee pirot
1
2
3
Habanero
1
2
3
157
Cultivar Plant no.Stem diameter (cm.)
SPAD chlorophyll reading
(SCMR)
T1 T2 T3 T1 T2 T3
Bhut jolokia
1
2
3
Table 3 Stem diameter and SPAD chlorophyll reading of 3 hot pepper cultivarstreated by 3 levels of drought stress
2.4 Assessment of resistance and tolerance
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3
Akanee pirot
1
2
3
Habanero
1
2
3
158
Chapter V: Plant and environmental stress
Chapter V: Plant and Abiotic Stress
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159
Chapter V: Plant and environment stress
5.1 The stress environment (Dr. Suchila)
5 3 Pl t b di d i ld t bilit
5.2 Responses of plant to environmental stress (Dr. Suchila)
Chapter V: Plant and environment stress
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5.3 Plant breeding and yield stability (Dr. Suchila)
5.5 Selection for yield and environment (Dr. Sanun)
5.6 Physiology and genetic implication (Dr. Sanun)
5.4 Models and applications (Dr. Sanun)
160
5.1) The Stress Environment
• The 1972 crisis (World imbalance of population and food supply) demonstrated how closely the world borders on severe food shortages The food production limitations
5.1 The stress environment
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shortages. The food production limitations other than man‐made stresses, such as air pollution, and pest losses are: (a) availability of productive soil, and (b) a climate supportive to crop growth (Christiansen, 1982)
161
Land availability• ~10% of the world’s arable may be categorized as free of stress
• ~20% of the land is under some kind of mineral stress
5.1 The stress environment
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• ~26% is affected by drought stress
• ~15% is affected by freezing stress
Source: Dudal (1976, quoted by Blum, 1986)
162
1
Climatic restriction
It has been estimated that 60‐80% of variability in crop production is a result of whether fluctuation (temperature and water availability) Well known and senior breeder in
5.1 The stress environment
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availability). Well known and senior breeder in USA concluded that modern technology has little influence on the susceptibility of non‐irrigated crops to whether (Christiansen, 1982).
163
Types of abiotic environmental stress
Temperature: heat, cold
Water: drought, water logging
5.1 The stress environment
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p
Salinity & Mineral
164
Stress environments:
Water stress
• Very little of the world’s water is used by plants. Nace (1969, quoted by Christiansen, 1982) estimated that only 2.5% of the world supply is non‐oceanic or fresh, and that only 0.007% of all water is used by
5.1 The stress environment
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y yplants.
• In rain‐fed agriculture the problem of short‐term (10‐20 day) drought is common and exacts an intangible and real toll on productivity. This incidence at critical
periods of cropping cycles can be disastrous.165
Stress Environment: Heat stress
• Most of the world’s agriculture is restricted by temperature to the confines of 50° South to 50° North latitudes (low temperature). High
temperature is a limiting factor in much of lowland tropics.
• The critical thermal regimes affect plant phenology, developmental phases, growth rates, yield components and final yield. Growth and development processes follow distinct temperature‐response
5.1 The stress environment
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development processes follow distinct temperature response curves, displaying a peak or a plateau at what are defined as an optimum temperature. Any temperature outside the optimum is non‐optimum, and may be defined as stress temperature. Various plant processes (different crops and growth stages) may have different temperature optima, or different threshold stress temperature (related to heat‐sums or degree‐days).
166
According to threshold temperatures: two annual crops groups:
Cool season vegetable
cabbage
cauliflower
Warm season vegetable
o Eggplant
o Corn
5.1 The stress environment
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sweet pea
melon
sweet pepper
Allium
o pumpkin
o Water melon
o Cucumber
o Yard long bean
167
Stress environment: salinity stress
• Soil properties that determine its mineral effects on plants are mineral content, organic matter content, CEC, permeability etc.
• The ultimate definition of a given soil‐nutritional problem, at the breeder’s level is best reached by growing and analyzing
5.1 The stress environment
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the given crop in the target environment.
• The complexity of the formation of saline soils is affected by rainfall, mineral weathering, fossil salts etc. It is important for the breeder to recognize two major and different saline environments (saline soil, and non‐saline soil irrigated with saline or brackish water).
168
1
• Avoidance: The environmental factor is excluded from the plant tissue (J. Lewitt, )
5.2) Responses of Plant to Abiotic Stress
Physiological standpoint:
Chapter V: Plant and environment stress
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• Tolerance: The environmental factor penetrates the tissue but tissue survives
• Resistance imply to Tolerance
169
Criterion for the development and use of
screening tests:
Generic variation should occur in germplasm
Screening for environmental stress
5.2 Responses of plant to abiotic stress
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heritability (h2) for the trait should be greater than h2 for yield
Trait should be correlated with yield
Screening test should be easy, rapid and economical to apply
170
Importance of resistant or tolerant variety
• The genetic improvement of stress resistance is an economically viable solution. On the other hand, stress‐resistant crop varietiesshould not necessary associated with
5.2 Responses of plant to abiotic stress
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ysituations of poor or marginal crop performance, or their importance should not be limited to conditions of low‐input and high‐risk farming (value added crop, nitch market;Blum, 1986).
171
5.3) Plant Breeding and Yield Stability
Chapter V: Plant and environment stress
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172
Drought Stress for Breeder standpoints
“Drought stress”:short term (duration) drought stress
5.3 Plant breeding and yield stability
5.3.1 Plant breeding for Environmental stress: “Drought stress”
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Drought stress is implied a total yield loss.
short-term (duration) drought stress
Drought stress occurs at any subpotential yield level. The improvement of yield at any of these levels is the main purpose of breeder.
173
“Drought is responsible for severe food shortages and famine in developing countries.”
Breeding for drought stress resistance
Drought stress is induced by inability of the
5.3 Plant breeding and yield stability
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Drought stress is induced by inability of the crop to meet its evapotranspiration demand. Therefore the reduction of the ETactual/ETmaximum
is interested. Potential evapotranspiration = PEActual evapotranspiration = ET
174
1
The goal of breeder: to obtain economic yield
Breeding for drought stress resistance
Yield = T x WUE x HI
5.3 Plant breeding and yield stability
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Where T= total seasonal crop transpiration
WUE = water use efficiency
HI = Crop harvest index
(Source: J.B. Passioura and C.T. de Witt)
175
Plant traits affecting drought response
Phenology: early or late
Plant development and size
Non‐senescence
Stem reserve utilization
Chapter V: Plant and Abiotic Stress
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Plant surface: wax
Root: length & dept
Photosynthetic systems
Water use efficiency
176
Mechanisms of drought resistance
• Escape: early maturity characteristic under optimum condition at the beginning season
• Avoidance: Decreasing water loss by extract soil moisture efficiency
Chapter 5: Plant and Abiotic Stress5.3 Plant breeding and yield stability
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• Tolerance: Some species more toleranat of post anthesis drought, and can produce appreciable yield under stress
• Recovery: The environmental factor is excluded some species are able to recover after short duration drought stress
177
5.3.1 Plant breeding for environmental stress: “Heat (high temperature) stress”
Different objective between the breeder inTropical vs temperate area
Screen under hot target environment (hot spot):
5.3 Plant breeding and yield stability
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Using controlled green house of growth chamber or phytotron room
g ( p )worldwide research stations of international seed Co.
178
5.3.1 Plant breeding for environmental stress: “Salinity Stress”
Soil salinity: ~ 95% million hectares worldwide
Soil salinity: the accumulation of dissolved salts in the soil
Different responses among different crops
5.3 Plant breeding and yield stability
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Tolerant Moderately tolerant
Moderately sensitive
sensitive
Cotton Sugar beet Peanut Strawberry
Barley Soybean Corn Potato
Sorghum Rice Tomato
Sweet potato Onion179
Management of resistant/tolerant varieties
The concept and role of genotype x environment interaction in plant breeding
5.3.2) Yield Stability
5.3 Plant breeding and yield stability
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Management of resistant/tolerant
varieties
180
1
The concept and role of genotype x environment (G x E) interaction in plant breeding
Plant breeder are interested in developing high yielding cultivars with sustained or stable high performance over seasons and years; that is “Yield stability”
5.3 Plant breeding and yield stability
123752: Suchila, KKU, Thailand
years; that is Yield stability .
G x E occurs when two or more genotypes are compared across different environments, and their relative performance are found to differ.
181
Stability is the ability of plant that can maintain yield under the environment variables.
5.3 Plant breeding and yield stability
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Different responses of chilli cultivars under two different location environment
182
1. Estimation of plant growth or characters in the different environments for many years
2. Study of the Regression coefficient between the output of each crop varieties with yield of all cultivars grown at that l ti (Fi l &
Models and application
5.3 Plant breeding and yield stability
123752: Suchila, KKU, Thailand
all cultivars grown at that location (Finlay & Wilkinson and Eberhart & Russel)
3. Measuring of the combining ability of varieties, that growing in different environments.
4. Determined by GGE biplot
183
GGE-Biplot
GGE‐biplot is an effective tool for:
Mega‐environment analysis (e.g. “which‐won‐where”
pattern), whereby specific genotypes can be recommended
to specific mega‐environments (Yan, 2003; Yan, 2006)
5.3 Plant breeding and yield stability
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to specific mega environments (Yan, 2003; Yan, 2006)
Genotype evaluation (the mean performance and stability)
Environmental evaluation (the power to discriminate
among genotypes in target environments)
184
GGE-Biplot
5.3 Plant breeding and yield stability
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In total phenotypic variation, E explains most of the variation and G and G x E are usually small (Yan, 2002)
185
Classification of G x E interaction
No G x E interaction: one genotype performs better than the others across all environments
A non cross over G x E interaction: one genotype outperforms the others across all environments, but that genotype may exceed the other genotype 20 unit in one
5.3 Plant breeding and yield stability
123752: Suchila, KKU, Thailand
g yp y g ypenvironment (env.) But 60 unit in the other env.
A crossover G x E interaction: (Most important to plant breeder)
a genotype A is more productive in one env., while the other genotype more productive in the other env.
Combined G x E interaction: one genotype increase the other genotype decrease 186
1
Finlay & Wilkinson and Eberhart & Russel
5.3 Plant breeding and yield stability
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187
Stability analysis
5.3 Plant breeding and yield stability
123752: Suchila, KKU, Thailand
188
Descriptions of environments where trials were conducted
during 2009–2011
Different altitudes temperatures
5.3 Plant breeding and yield stability
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Different altitudes, temperatures, RH%, rainfalls, solar radiations and soil properties
(6 environments)
189
Combined analysis of variance for yield and capsaicinoid traits of six
chili cultivars evaluated in six environments during 2009–2011
5.3 Plant breeding and yield stability
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Interactions between cultivars and environments (C x E) was found in all traits studied.
190
Stability analyses for yield and capsaicinoids of six chili cultivars
grown at six environments during 2009-2011
Dallay Khorsaney performed combined G x E inteaction
5.3 Plant breeding and yield stability
123752: Suchila, KKU, Thailand
KKU-P-21041 performed stable fruit yield across all environments
191
GGE-Biplot
GGE biplot for dry fruit yield (A) and capsaicinoids content (B) of six cultivars across six environments. Environment
identification, KK1: Khon Kaen rainy season 2009; CM1: Chiang Mai rainy season 2009; LB: Lobesa rainy season
2010; KB: Kabesa rainy season 2010; KK2: Khon Kaen dry season 2010-2011; CM2: Chiang Mai dry season
2010 -2011.
5.3 Plant breeding and yield stability
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GxE interactions complicate the process of selecting of genotypes with superior performance.192
1
Gerplasm
Chapter VI: Management of resistance /tolerance varieties
123752: Suchila, KKU, Thailand
Breeding Variety release
193
Remarks:
Breeder should note that resistance or tolerance
is not durable.
Chapter VI: Management of R & T var.
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“The resistant/tolerant varieties need to be improved time to time”i.e. genes pyramiding or multi-lines improvement.
194
Breeder seed
Foundationseed
breeder and institute
breeder and institute
Seed production of resistant/tolerant varieties(similar protocol to the normal seed production)
Chapter VI: Management of R & T var.
123752: Suchila, KKU, Thailand
Certified seed
Registered seed
grower or government or private sectors
seed registered officer
195
Variety registration & release
International procedure
(see CSSA guild lines)
Search and discuss about:
Chapter VI: Management of R & T var.
123752: Suchila, KKU, Thailand
Thailand procedure(see www.doa.go.th/pvp/)
196
Registration procedureInternational procedure(see CSSA guild lines)
• Cultivar name• Scientific name• Designation during
Thailand procedure(see www.doa.go.th/pvp/
• Cultivar name• Scientific name• Name of organizations
Chapter VI: Management of R & T var.
123752: Suchila, KKU, Thailand
g gdevelopment
• Name of organizations or institutions
• Brief description of the cultivars
• etc.
gor institutions
• Brief description of the cultivars
• Information for cultivar development
• etc.197
www.doa.go.th/pvp/
123752: Suchila, KKU, Thailand
Registered for PVP in Thailand
Department of Agriculture, Ministry of Agriculture and cooperatives 198
1
Patents: USA vs Thai
Chapter VI: Management of R & T var.
123752: Suchila, KKU, Thailand
http://www.google.com/patents/US20090019600
199
Acquaah, G. 2007. Principles of Plant Genetics and Breeding. Blackwell Publishing.
Blum, A. 1988. Plant breeding for stress environments. CRC Press, Inc. Florida.
www.sciencecampaign.org.uk/.../2009/beddingtonpresentat...
Key References
123752: Suchila, KKU, Thailand
http://www.agnet.org/library.php?func=view&style=type&id=20120104150721
http://www.google.com/Adaptation and yield stability.htm
http://www.google.com/Patent WO2009098685A2 ‐ Disease resistant pepper plants
http://plantandsoil.unl.edu/croptechnology2005/pages/animationOut.cgi?anim_name=RootUptake.swf
http://www.google.com/patents/US8642845200