Citation: Pande, S., Singh, G., Narayana Rao, J., Bakr, M.A., Chaurasia, P.C.P., Joshi, S., Johansen, C., Singh, S.D.,Kumar, J., Rahman, M.M., and Gowda, C.L.L. 2001. Integrated management of botrytis gray mold of chickpea. (InEn. summaries in En, Fr.). Information Bulletin no. 61. Patancheru 502 324, Andhra Pradesh, India. InternationalCrops Research Institute for the Semi-Arid Tropics. 32pp. ISBN 92-9066-434-7 Order Code IBE 061.

AbstractBotrytis gray mold (BGM) caused by Botrytis cinerea is an important disease of chickpea worldwide. Its origin, distribution,losses, symptoms, causal organism and its variability, epidemiology, and host range are briefly reviewed. Information onhistopathology and host-pathogen interaction with respect to chickpea as a host is not available. Therefore, efforts weremade to supplement this part of the literature from other hosts of B. cinerea. Attempts have also been made to assemblethe information on integrated disease management (IDM) of BGM. The IDM components reviewed are host-plantresistance, agronomic and cultural practices including effects of sowing date (escape), row spacing, plant type, andintercropping; management by chemicals which include seed treatment and foliar sprays; and management with biologicalagents. Integrated management of BGM in chickpea involves use of BGM-resistant cultivars with improved agronomicand cultural practices including economical use of fungicides, but these practices are not yet sufficiently refined to beadapted by resource-poor farmers. Therefore, farmers’ participatory on-farm research is needed to devise appropriatepackages of these strategies for BGM endemic areas.

RésuméLa pourriture grise (BGM) causée par Botrytis cinerea est une importante maladie qui affecte le pois chiche dans lemonde entier. L’origine de cette maladie, sa distribution, les pertes qu’elle cause, ses symptômes, l’agent causal, savariabilité, son épidémiologie et l’éventail des hôtes sont brièvement examinés. Les informations sur l’hystopathologieet l’interaction hôte-pathogène pour le pois chiche en tant que plante hôte ne sont disponibles. Des efforts ontdonc été faits pour compléter cette partie de la littérature à partir d’autres hôtes de B. cinerea. Des tentatives ontégalement été faites pour réunir des informations sur la lutte intégrée (LI) contre le BGM. Les composantes de laLI qui ont été examinées sont : la résistance de la plante hôte; les pratiques agronomiques et culturales dont leseffets de la date de semis (décalage) ; l’espacement des rangs ; le type de plantes et les cultures intercalaires ; lalutte à l’aide des produits chimiques, notamment le traitement des semences, les pulvérisations des feuilles et lalutte à l’aide d’agents biologiques. La lutte intégrée contre le BGM chez le pois chiche nécessite l’utilisation decultivars résistants au BGM, accompagnée par des pratiques agronomiques et culturales améliorées, y comprisl’utilisation de fongicides. Cependant, ces pratiques ne sont pas suffisamment affinées pour être adoptées par despaysans pauvres. Par conséquent, la recherche participative en milieu paysan est nécessaire pour concevoir despaquets appropriés de ces stratégies pour les zones où le BGM est endémique

ACKNOWLEDGEMENTSThe authors are indebted to all the scientists and researchers who contributed to the botrytis gray mold (BGM)research worldwide. Our specific grateful thanks to investigators of BGM of chickpea in Bangladesh, Nepal andIndia whose research findings on various aspects of BGM and its management laid a solid foundation for thiscompilation. Contribution to word processing of original manuscript and subsequent corrections and the coordinatingrole played by Ms I Radha is sincerely acknowledged. We appreciat the technical editing by Ms Rita Banerjee formaking this information bulletin reader-friendly. The coordination of the production process by Mr. T R Kapoor isgratefully acknowledged.

Cover: Integrated management of botrytis gray mold of chickpea in a farmer’s field in Nepal. Top: a healthy cropraised with disease management practices (inset: healthy plants with abundant pods and leaves); bottom: a diseasedcrop grown without disease management practices (inset: diseased plants with dead shoots and flowers).

The research activities were supported by the Asian Development Bank (ADB)and donors supporting ICRISAT’s unrestricted core activities.

Information Bulletin No. 61

S Pande, G Singh, J Narayana Rao, M A Bakr, P C P Chaurasia,

S Joshi, C Johansen, S D Singh, J Kumar, M M Rahman,

and C L L Gowda

International Crops Research Institute for the Semi-Arid TropicsPatancheru 502 324, Andhra Pradesh, India.

2001

Integrated Management ofBotrytis Gray Mold of Chickpea

ICRISAT

About the Authors

S Pande Principal Scientist (Pathology), Natural Resource Management Program, ICRISAT,Patancheru 502 324, Andhra Pradesh, India.

G Singh Senior Plant Pathologist (Pulses) and Head, Pulses Program, Punjab AgriculturalUniversity, Ludhiana 141 004, Punjab, India.

J Narayana Rao Scientific Officer (Pathology), Natural Resource Management Program,ICRISAT, Patancheru 502 324, Andhra Pradesh, India.

M A Bakr Senior Scientific Officer, Plant Pathology Division, Bangladesh Agricultural ResearchInstitute (BARI), Joydebpur, Gazipur 1701, Bangladesh.

P C P Chaurasia Senior Scientist, Plant Pathology, Regional Agricultural Research Station,Nepal Agricultural Research Council (NARC), Tarahara, Nepal.

S Joshi Senior Scientist, Plant Pathology Division, Nepal Agricultural Research Council(NARC), Khumaltar, P O Box 5459, Kathmandu, Nepal.

C Johansen Consultant, Apartment 2-B, Palmdale, Plot No. 6, Road No. 104, Gulshan-2,Dhaka, Bangladesh.

S D Singh Senior Scientist (Pathology), Genetic Resources and Enhancement Program,ICRISAT, Patancheru 502 324, Andhra Pradesh, India.

J Kumar Principal Scientist (Breeding), Genetic Resources and Enhancement Program,ICRISAT, Patancheru 502 324, Andhra Pradesh, India.

M M Rahman Project Director, Pulses Research Center, Bangladesh Agricultural ResearchInstitute (BARI), Ishurdi, 6620, Pabna, Bangladesh.

C L L Gowda Principal Scientist (CLAN), Genetic Resources and Enhancement Program,ICRISAT, Patancheru 502 324, Andhra Pradesh, India.

The designations employed and the presentation of the material in this publication do notimply the expression of any opinion whatsoever on the part of ICRISAT concerning the legalstatus of any country, territory, city, or area, or of its authorities or concerning the delimitationof its frontiers or boundaries. Where trade names are used, this does not constitute endorsementof or discrimination against any product by the institute.

Preface vIntroduction 1Geographical distribution and economic importance 3Disease symptoms 3Causal organism 5Variability 6Epidemiology and biology 6Survival in diseased crop debris and soil 6Seed transmission 6Collateral host 7Seasonal disease development 7Disease management 8Agronomic and cultural management 8

Sowing date 8Row spacing 9Plant type 9Intercropping 10

Management using host-plant resistance 11Screening techniques 11

Growth room screening technique 11Growth chamber screening technique 12Cut-twig screening technique 12Field screening technique 12

Identification of resistance 12Breeding for resistance 13Wide hybridization 14Intraspecific hybridization 14

Management by chemicals 16Seed treatment 16Foliar spray 16

Management using biological agents 18Integrated control 19Conclusion 19Future prospects – Use of transgenics 20References 21

Contents

Preface

Chickpea (Cicer arietinum L.) is one of the most important grain legume crops. It is widelyconsumed in South Asia as dhal and is a very important source of protein in the diets ofresource-poor farm families. However, farmers in this region face many biotic and abioticconstraints to successful cultivation of the crop. Among these, the fungal disease botrytis graymold (Botrytis cinerea Pers. ex Fr.) (BGM) is one of the most serious problems, especially inNepal, India, Bangladesh and Pakistan.

ICRISAT with partner institutes from the above countries has been working for the past 10years to develop sustainable management strategies for BGM. Although initial studies focusedon judicious use of fungicides and host-plant resistance, it soon became clear that the leastrisky solution was to develop an integrated management strategy based on a number ofcompatible and reinforcing actions. These include host-plant resistance, foliar fungicide sprays,seed treatment, sowing date (escape), plant density, intercropping, and biological control.

This bulletin reviews and summarizes the progress to-date in developing and testing, throughon-farm participatory research, appropriate integrated management packages for BGM inendemic areas. It is clear that substantial progress has been made in developing the package,but further work is needed to ensure uptake by farmers. This will be accomplished in somecountries, e.g., Nepal, through new projects initiated recently. At the same time, additionalmanagement strategies are also being studied, e.g., transgenic approaches that may in futurebe integrated with the existing methods.

The authors should be congratulated for their committed efforts to tackle this seriousdisease problem, and encouraged to continue their endeavors in disseminating the most effectivemanagement packages to farmers in BGM-endemic areas of South Asia.

Jill LennDeputy Director General - Research

ICRISAT

1

Introduction

Botrytis gray mold (BGM), caused byBotrytis cinerea Pers. ex Fr., is an importantdisease of chickpea (Cicer arietinum L.) inIndia, Bangladesh, Nepal, and Pakistan. Thedisease has also been reported as a majoryield reducer in Australia and Argentina. Itcan cause substantial yield losses (Fig. 1) ifthe conditions are favorable during vegetativeand reproductive growth stages of the crop(Singh et al. 1982, Grewal and Laha 1983).Use of cultivars resistant to BGM is the bestway to control the disease. But, stablesources of host-plant resistance to BGM arenot available in the germplasm and breedingmaterial (Laha and Khatua 1988, Ahmad

1989, Singh et al. 1992). However, high levelsof resistance to BGM have been reported inwild Cicer species (Singh et al. 1982, 1991,1992; Haware et al. 1992). Several seeddressing and foliar fungicides have been foundeffective (Grewal and Laha 1983, Singh andBhan 1986b, Singh and Kaur 1990) but theuse of fungicides to manage the disease hasnot been widely adopted by resource-poorfarmers (Singh and Bhan 1986b). Hence,integrated management of BGM usingavailable levels of host-plant resistance,improved agronomical practices, increaseddependence on biological agents (if available),and targeted spraying with effectivefungicides is being advocated (Pande et al.1998, 2000).

Figure 1. Chickpea crop loss due to botrytis gray mold.

2

Figure 2. Geographical distibution map of Botrytis cinerea, showing countries where it occures on chichpea.

Figure 3. Areas in the Indo-Gangetic plain (IGP) countries where Botrytis gray mold (BGM) of Chickpeais endamic.

3

Geographical Distributionand Economic Importance

The occurrence of BGM on chickpea wasfirst reported by Shaw and Ajrekar in 1915,but Carranza (1965) reported its first fieldincidence in Argentina. The disease has beenreported from Argentina, Australia,Bangladesh, Canada, Chile, Colombia,India, Myanmar, Nepal, Pakistan, Spain,Turkey and USA (Malik et al. 1993, Neneet al. 1984, Haware and McDonald 1992,1993). Its geographical distribution isillustrated in Fig.2.

There are several reports of serious lossescaused by BGM. Carranza (1965) reporteda 95% crop loss in Jujuy province ofArgentina due to BGM. The disease wasresponsible for heavy losses in the Indo-Gangetic plains of India during 1979 82(Grewal and Laha 1983). During 19781979 an epidemic of BGM destroyedchickpea crop completely over an area of20,000 ha in the states of Punjab, Haryana,Uttar Pradesh and Bihar, India (Singh et al.1982, Grewal and Laha 1983, Grewal1988). In Bangladesh, the damage causedby the disease was estimated to be 80 90%in 1988 and 70 80% in 1989 (Bakr andAhmed 1992). In Nepal, the disease occursalmost every year and an estimated loss of66% in the experimental fields and about15% in farmers’ fields was reported (Joshi1992). Recently, more than 80% crop lossesdue to BGM have been observed in the Indo-Gangetic plains of Bangladesh, Nepal, andnorth-western India (Fig. 3; Pande 1998).

Disease SymptomsThe pathogen infects all aerial parts of theplant (Figs. 4 and 5), flowers being the most

Figure 4. Tender twigs showing symptoms with graymass caused by botrytis gray mold.

Figure 5. Flowers showing rotting and gray masscaused by botrytis gray mold.

4

Figure 6. Symptoms of botrytis gray mold under field conditions.

vulnerable (Joshi and Singh 1969,Kharbanda and Bernier 1979, Haware andNene 1982, Laha and Grewal 1983b).Heavy mortality of flowers results in poorpod formation. Under natural conditions,the disease first appears in isolated patcheswhere the crop has dense plant canopy whenhigh relative humidity (≥95% RH) and lowtemperature (20 25 C) prevails (Fig. 6).The symptoms appear on chickpea stems,branches, leaves, flowers, and pods as grayor dark brown lesions covered with erecthairy sporophores and masses of hyalinespores (Figs. 7 and 8). The fungal growth is

evident on all plant parts hidden under denseplant canopy, particularly on flowers andpods. Stem lesions are 10 30 mm long andgirdle the stem completely. Affected leavesand flowers turn into a rotting mass. Onthick, hard stems, the gray mold growth isgradually transformed into a dirty gray masscontaining dark green to black sporodochia.Sometimes, tiny black sclerotia can be seenon dead tissues. The sclerotia are small, darkbodies and should not be confused with largerblack or dark brown sclerotia embedded inwhite mycelium of Sclerotinia sclerotiorum(Lib.) de Bary (Joshi and Singh 1969).

Figure 8. Dark brown lesions on pods caused bybotrytis gray mold.

Figure 7. Stem showing grayish fungal sporulationcaused by botrytis gray mold.

5

When BGM affects pods, no seeds or onlysmall, shrivelled seeds are formed.Occasionally, grayish-white mycelia may beseen on immature seeds. Lesions on the podare water-soaked and irregular, sometimes withblack sclerotial bodies scattered in the infectedareas.

Causal Organism

The genus Botrytis was first erected byMicheli (1929), and since then it hasbecome widely known as a group of fungicausing economically and potentiallyimportant plant diseases. This is particularlytrue of those forms which group togetheras the form species Botrytis cinerea. Thesespecies tend to be concentrated in thetemperate regions between 25 and 30latitudes, where they occur on a variety ofcrop plants.

The causal organism of BGM of chickpeais B. cinerea. and its teleomorph isBotryotinia fuckeliana (de Bary) Whetzel(Grooves and Loveland 1953). The

Figure 9. Growth of Botrytis cineria on potatodextrose agar.

Figure 10. Mycelium of Botrytis cinerea showingclusters of conidia.

Figure 11. Conidiophore of Botrytis cineria bearingsterigmata and conidia.

teleomorphic state of this fungus has beenproduced from sclerotia of B. cinereainfecting chickpea in India (Singh 1997).

Botrytis cinerea is a necrotrophic funguswell known for its extensive host range, widedistribution globally, extreme variability andadaptability to a wide range of environmentalconditions. The fungus grows profusely ondead flowers and potato dextrose agar(Fig. 9). Initially, growth of the fungus is whiteand cottony, turning light gray with age (Ellisand Waller 1974). The fungal mycelium isseptate and brown, and young hyphae are thin,hyaline, and 8 16 µm wide. The conidiophoresare light brown, septate, erect, with tips slightlyenlarged bearing small pointed sterigmata(Figs. 10 and 11). The conidia are hyaline, one-celled, oval or globose, and are borne in

6

clusters on short sterigmata (Fig.12). Conidiafrom BGM lesions on chickpea plants measure4 25 × 4 18 µm and from potato dextroseagar 4 16 × 4 10 µm. The sporodochia onthe host surface are 0.5 5.0 µm in diameter(Joshi and Singh 1969). The culturalcharacteristics and sporulation of this fungusvary with synthetic medium, temperature andother ecological factors (Coley-Smith 1980,Laha and Grewal 1983a).

Variability

There have been very few studies onvariability of B. cinerea but the pathogenproduces different pathotypes or races tocope with host diversity and fluctuatingenvironmental conditions. Joshi and Singh(1969) and Singh (1970) observed theformation of sclerotial and/or sporodochialbodies on chickpea plants infected with B.cinerea in the Tarai region of Nainital, India.But Pandey (1988) did not observe thesebodies in the same area. This indicates thatcertain isolates of B. cinerea are capable ofproducing sclerotia while others do not, andwithin a location there may be more thanone isolate of the fungus. Singh and Bhan(1986a) identified four physiological racesof the pathogen from northern India, andRewal and Grewal (1989b) reported fivepathotypes of B. cinerea from India.

Epidemiology and Biology

Survival in diseased cropdebris and soil

Botrytis cinerea survives in the soil in theform of mycelia and sclerotia from one cropseason to the next. Infested soil and infected

plant debris are the main sources of primaryinoculum (Mahmood and Sinha 1990).Madhu Meeta et al. (1986b) reported thatB. cinerea survived in plant debris for 180days in the soil at a depth of 6 cm. The funguswas found to be viable in infected seed andplant debris stored at 18 C for 5 years (Grewal1988). But, Singh (1989) found that B. cinereadid not survive beyond 8 months in chickpeadebris at a depth of 10 cm in the soil, and thatthe survival and recovery of the fungus wasmore at 5 10 C. Singh and Tripathi (1992,1993) observed the survival of B. cinerea ininfected chickpea debris at a depth of 25 cmfor 8 months, i.e., until the following season.

Seed transmission

The seedborne nature of B. cinerea in chickpeahas been reported by many workers (Cother1977, Laha and Grewal 1983b, Haware etal. 1986, Sandhu and Sah 1988, Burgess etal. 1997a). The fungus remained viable in theinfected seed until the next growing season(Madhu Meeta et al. 1986b). Grewal (1988)reported that the fungus was externally andinternally seedborne. Singh (1989) recordedits survival in chickpea seed for 8 months.He further recorded that the survival of this

Figure 12. Conidia of Botrytis cinerea on potatodextrose agar.

7

fungus was affected by ambient temperatureduring storage of the seed. Singh and Tripathi(1992) indicated that the survival andrecovery of the fungus was significantly moreat 5 10 C. They also reported that thesurvival period decreased with the increasein storage period and the pathogen did notsurvive in the seed at 40 C. Despite the factthat infected seeds carry the inoculum for aconsiderable period, the role of seedborneinoculum is not yet established as a source ofprimary infection in the field.

Collateral host

Botrytis cinerea is a facultative parasite andhas a wide host range. It causes gray molddisease in a number of crop plants, such asstrawberry, grapevine, apple, cabbage,carrot, cucumber, eggplant, lettuce, pepper,squash, tomato, and several ornamentalssuch as chrysanthemum, dahlia, lilly, roses,gladiolus, and tulips. Madhu Meeta et al.(1988) reported that B. cinerea could infectpea (Pisum sativum L.) and several weedsunder greenhouse conditions. Rathi andTripathi (1991) also observed that it couldinfect eight cultivated species and 21 weeds.Singh (1997) reported that B. cinerea couldinfect about 100 plant species, includingornamental plants, vegetables, fruit crops,field crops, and several weeds.

Seasonal diseasedevelopment

An understanding of the environmentalfactors that contribute to disease initiationand epidemic development are necessary torecommend control methods. Theimportance of sclerotia and conidia in

epidemic initiation and build up may vary,depending upon regions. In some areas, initialinfections may be through soilbornepropagules. If conditions are favourable forconidial production, these early infectionslead to an increased inoculum load of conidialater in the season, and this results in anincreased severity of BGM. As the fungus alsosurvives on infected chickpea seeds,seedborne inoculum plays an important rolein BGM initiation and further diseasedevelopment in Australia (Cother 1977).Since B. cinerea infects a very wide range ofplants, some of which are perennials, the roleof collateral hosts in the survival of the fungusand infection of chickpea from one seasonto another can also influence epidemicdevelopment. The relative epidemiologicalimportance of seedborne inoculum and othersources of inoculum such as plant debris hasnot been fully elucidated for gray mold ofchickpea (Haware and McDonald 1993). Itis, therefore, difficult to predict the likelihoodof its epiphytotics. Further work on theepidemiology of BGM in chickpea is clearlywarranted.

The important environmental factorsthat contribute to BGM development arerelative humidity, duration of leaf wetness(Butler 1993), and temperature. Flow of airmodify humidity and temperature in cropcanopies, and hence, BGM developmentdepends on the density of the crop canopy.Bakr et al. (1993) reported that theincidence of the disease was high when themean temperature ranged from 17 to 28 Cand relative humidity from 70 to 97%.Singh and Kapoor (1984) observed thatthere was an increase in the diseaseintensity with increase in the leaf wetnessperiod beyond 12 h day-1. Rewal andGrewal (1989a) reported that 5 C was the

8

minimum, 20 C the optimum, and 30 Cthe maximum for conidial germination ofall three isolates studied, and also that allthe three isolates required differential lightintensities and relative humidity forconidial germination. The fungus couldgrow in a wide range of temperatures, butthe optimal temperature for fungal growthand sporulation was 25 C (Mahmood andSinha 1990). A relative humidity of 95%or above for a few hours during the dayand a dense canopy were most favorablefor infection and rapid spread of the disease(Tripathi and Rathi 1992).

Disease Management

Effective management of BGM is veryimportant, since the disease causes heavydamage to chickpea. Since a high level ofresistance to BGM is not available in thecultivated chickpea, it is necessary to combineavailable resistance levels with othermanagement options for minimizing thedisease. Although repeated application offungicides can control BGM, it may not bepracticable for resource-poor farmers inBGM-endemic areas of South Asia and itcould also result in the production of newand/or resistant strains of the fungus (Bhanand Chatrath 1994). Judicious use offungicides as a seed treatment and/or foliarspray in an integrated disease managementsystem could be very economical andaffordable to the resource-poor farmer.Disease management systems integrating theuse of available levels of host-plant resistance,agronomical and cultural, chemical, andbiological disease control components inseveral combinations, need to be developed,evaluated, and promoted. Components ofBGM management are as follows.

Agronomic and culturalmanagement

Agronomic and cultural management ofBGM in recent years has been very welldemonstrated in India, Bangladesh, andNepal. Early sowings, high seed rates, closerrow spacings, and traditional use of bushygenotypes result in dense canopies, whichfavor development of BGM in chickpea.Thus, techniques which minimize canopydensity may alleviate disease intensity.

Sowing date

A crop sown after the optimal date ofplanting often becomes productive andavoids the most vulnerable growth stage andweather conditions favorable for BGMdevelopment. Early sowing results inexcessive vegetative growth due to moreresidual moisture and low to moderatetemperatures causing more BGM incidence,whereas late sowing reduces vegetativegrowth and hence lowers disease incidence.In both these situations, however, the grainyields were low (Brinsmead 1992, Hawareand McDonald 1992). Karki et al. (1989,1993) observed that delayed sowingsreduced BGM incidence but resulted in lowgrain yields. Further, they noticed positivecorrelations between plant height and BGMincidence. Haware and McDonald (1993)reported that delayed sowings reduced BGMincidence even in susceptible cultivars, butsignificantly reduced the grain yields(Table 1). Singh (1997) also observed thatthe late sown crop (around 20 Nov) inPunjab, India, showed significantly lowincidence of BGM. Bakr et al. (1997, 1998)also observed that early sown crops hadmore BGM both in compact and bushygenotypes than the late sown crops in

9

Bangladesh. Similar results were alsoobtained in India by Haware et al. (1997).

Row spacingWider spacings between rows reduce BGMincidence. This may be because of moreaeration in the crop canopy, which reducesperiods of leaf wetness to levels below therequirement for disease development. Reddyet al. (1988) observed that wider spacingsbetween rows reduced the RH in the cropcanopy and thereby caused lower incidenceof BGM in Nepal. Reddy et al. (1993)recorded similar results at Pantnagar, India.Haware and McDonald (1992, 1993)reported that BGM incidence was lower atwider row spacings than at closer spacingsin both fungicide sprayed and unsprayedplots. Bakr et al. (1993) reported that widerspacings alone reduced BGM disease

significantly in Bangladesh. Haware et al.(1997 see Table 2), Bakr et al. (1997), andShrestha et al. (1997) also reported thatwider row spacings reduced BGM diseaseincidence and resulted in higher grain yields.

There are not many studies on the effectof sowing geometry, row spacing, and roworientation, with respect to natural winddirection, and their effects on the leafwetness and disease incidence. Informationon these aspects would be useful to furtherimprove the agronomic management of thisdisease.

Plant typeErect and compact growth habit genotypesshow lower BGM incidence than bushy andspreading genotypes in India (Reddy et al.1988, 1990, 1993; Haware and McDonald1992, 1993; Haware et al. 1997), Nepal

Table 1. Influence of date of sowing (DS) of chickpea cultivars on gray mold severity1 and grain yield1,Pantnagar, India.

DS I2 DS II2 DS III2 DS IV2

Disease Plot yield Disease Plot yield Disease Plot yield Disease Plot yieldCultivars rating3 (kg)4 rating3 (kg)4 rating3 (kg)4 rating3 (kg)4

H 208 7.7 0.9 6.7 1.1 4.7 1.0 3.7 1.0Pant G 114 6.3 0.9 5.7 1.0 3.3 0.9 3.3 1.3K 850 6.0 1.0 6.0 1.1 4.3 0.9 3.3 1.3ICCV 88510 5.3 1.2 5.0 1.0 4.0 0.7 2.3 1.2ICCL 87322 4.7 1.2 4.7 1.1 3.3 0.9 2.3 1.1

SE

Disease rating Plot yield (kg)

Cultivar ±0.17** ±0.09Sowing ±0.15** ±0.08**

Cultivar × sowing ±0.34 ±0.17CV (%) 13 32

1. Mean of three replications.2. Dates of sowing; DS I = 31 Oct 1992; DS II = 14 Nov 1992; DS III = 29 Nov 1992; DS IV = 14 Dec 1992.3. Disease rating on a 1 9 point scale, where 1 = no infection and 9 = all plants killed.4. Plot size 2 m × 4 m** Significant at 1%.

Source: Haware and McDonald 1992.

10

(Shrestha et al. 1997) and Bangladesh(Bakr et al. 1997). An erect and compactgrowth habit is believed to improve aerationin the crop canopy thus making themicroclimate unfavorable for BGMdevelopment. However, no data are availableto verify this effect of plant type onmicroclimate under field conditions.

Intercropping

Little work has been done on the effect ofintercropping on BGM development inchickpea. Reddy et al. (1990) reported thatwhen chickpea was intercropped with linseedin Nepal, a marginal increase in chickpeagrain yield was obtained because of low BGMincidence and the linseed yield was a bonus.

Table 2. Effect of row spacing and Ronilan® sprays on the severity of botrytis gray mold, and grain yieldin chickpea, Pantnagar, India, 1992/93.

Disease rating1 Yield1

Treatment Cultivar Spacing (1-9 scale) (kg ha-1)

Sprayed2 ICCL 87322 30 × 10 4.3 119260 × 10 3.3 129145:15:453 4.3 120060:40:603 3.3 1176

H 208 30 × 10 5.7 126560 × 10 4.7 113945:15:45 4.7 115760:40:60 4.7 1086

Nonsprayed ICCL 87322 30 × 10 5.7 98960 × 10 4.3 98945:15:45 4.3 91160:40:60 4.0 1044

H 208 30 × 10 8.0 45960 × 10 6.3 34945:15:45 6.3 39260:40:60 6.0 399

SECultivars ±0.12** ±85.1Spacings ±0.17** ±120.4Sprayings ±0.12** ±85.1**Spacing × sprayings ±0.25 ±170.2Cultivar × spacing ±0.24 ±170.2Cultivar × spraying ±0.17* ±120.4*Cultivar × spraying × spacing ±0.35 ±240.7

CV (%) 12.1 45.4

1. Mean of three replications.2. Fungicide Ronilan (0.2%).3. Paired rows.** Significant at 1%; *Significant at 5%.

Source: Haware et al. (1997).

11

However, Karki et al. (1993) reported thatthere was no significant difference in BGMincidence in sole or intercrop with mustardin Nepal. Bakr et al. (1993) observed thatchickpea intercropped with linseed showeda reduced incidence of BGM in Bangladesh.These workers also suggested the use oflinseed as intercrop in the integratedmanagement of BGM disease of chickpea.

Management using host-plant resistance

Sowing disease-resistant cultivars is thecheapest and most effective method tocombat BGM, but adequately high levels ofresistance are not available in germplasmand breeding material of chickpea. However,high levels of resistance are available in wildCicer species. Hence, attempts have beenmade to identify stable resistance from wild

Cicer species using reliable screeningtechniques. The following techniques arebeing used to screen for BGM resistance incultivated and wild Cicer species and theirsegregating breeding material.

Screening techniquesTechniques to screen chickpea germplasm andbreeding material for BGM resistance havebeen developed by different research institutes.

Growth room screening technique. Thistechnique was developed at ICRISAT-Patancheru (Fig. 13). Seedlings of testgenotypes, along with a susceptible controlH 208, are raised in 10-cm diameter plasticpots (5 seedlings pot-1) filled with sand andvermiculite (4:1) mixture in a greenhouse.The pathogen, B. cinerea, is multiplied onpotato dextrose broth and incubated at 25 Cwith 12 h light and 12 h darkness in a

Figure 13. Growth room screening technique for botrytis gray mold resistance.

12

Percival incubator. Ten-day-old seedlings aretransferred to plant growth room andinoculated with 10-day-old conidial suspension(50 000 conidia mL-1) of the pathogen. Thegrowth room is maintained at 24±2 C and95 100% RH with 12 h light and 12 h darknesstill the end of the experiment. Diseasesymptoms start appearing 24 h afterinoculation. BGM severity is scored on 1 9scale (1 = no infection and 9 = all plants killed)3 days after inoculation and subsequently everyalternate day for 9 days.

Growth chamber screening technique. Thisscreening technique was developed by PunjabAgricultural University (PAU), Ludhiana,Punjab, India. The test lines are planted inpolythene bags (15 cm × 10 cm) containingsandy-loam soil in a greenhouse. Twenty-five-day-old plants along with susceptible check (G543 or H 208) are transferred to a growthchamber having controlled environmentalconditions (approximately 20 C, >90% RH,and alternate light and dark period). Afterwatering the pots, plants are inoculated byspraying spore suspension (50 000 sporesmL-1) of B. cinerea and enclosed for 6 dayswithin a moist chamber prepared withpolythene sheets supported by iron frames(46 cm × 46 cm). During incubation period,8 h light and 16 h dark periods are provided.The disease symptoms appeared after 24 h,and 100% mortality of susceptible lines andcontrol are recorded in 6 days after inoculation(Singh et al. 1982).

Cut-twig screening technique. Thistechnique consists of cutting 10 15 cm longtender shoots of chickpea plants with a sharpedged blade in the evening. The lowerportion of the twig is wrapped with a cottonplug and transferred to a test tube (150 mm× 10 mm) containing fresh tap water. The

tubes are placed in a test tube stand andinoculated along with twigs of susceptiblecheck (G 543 or H 208 or L 550).Inoculation method and post-inoculationincubation conditions are similar to thoseused for growth chamber-screeningtechnique. The disease symptoms appear24 h after inoculation and 100% mortalityof susceptible lines occurs 6 days afterinoculation (Singh et al. 1998).

Field screening technique. This techniqueis very efficient for large-scale screening ofgermplasm and breeding material insegregating generations (Fig. 14). The testlines are sown in 2 3 m long rows spaced at30 cm × 10 cm. Indicator-cum-infector rowsof a susceptible cultivar H-208 are sownafter every two test rows. When the plantsare 70 80-day old (at the onset of flowering),the field is irrigated in the morning and plantsare inoculated by spraying a spore suspension(50 000 spores mL-1) of 10-day old cultureof B. cinerea. From the morning of thefollowing day, sprinkler irrigation or perfo-spray system is run every day for about 15minutes after every 1 or 2 h from 0900 to1700 h depending upon the environmentalconditions. The inoculation of plants arerepeated twice at 10-day interval after thefirst inoculation. The disease severity isscored on 1 9 rating scale in mid-February,and first and last weeks of March.

Identification of resistanceExtensive screening (both in controlledenvironment and field screening at hot-spotlocations) of germplasm indicated that therewere no genotypes with high levels ofresistance (Haware and Nene 1982, Rathiet al. 1984). Rewal and Grewal (1989b)tested a large number of chickpea lines forBGM resistance and found ICC 1069 and

13

ICC 5035 to be relatively resistant. Laha andKhatua (1988) also reported that ICC 1069was most resistant to BGM in theirevaluation. Singh and Kaur (1989) reportedthat GL 84195 and GL 84212 were mostresistant and GL 86094 was moderatelyresistant to BGM. Ahmad (1989) foundonly moderately resistant cultivars to BGMin his screening in Bihar state of India.Several lines found resistant at Pantnagarwere susceptible at Rampur, Nepal (Hawareand McDonald 1993). Recently, chickpeagenotype ICCL 87322, a tall and erect type,was found to be resistant (BGM score ≤5)at hot-spot locations in India, (Haware etal. 1997), Bangladesh (Bakr et al. 1998) andNepal (P C P Chaurasia, Tarahara,unpublished). However, lines found resistantat one location can become susceptible atother locations because of the pathogenvariability (Singh and Bhan 1986a). Ashigher levels of resistance to BGM are not

available in cultivated chickpea, wild Cicerspecies have been screened under controlledenvironment to identify resistant sources.Cicer judaicum 182, C. judaicum ILWC19-2, and C. pinnatifidum 188 were foundto be highly resistant to BGM (Singh et al.1991, Haware et al. 1992).

Breeding for resistance

BGM-tolerant and resistant lines fromdifferent locations have been crossed withdesi and kabuli types in India (at ICRISAT,Pantnagar, and PAU) and in Bangladesh(Ishurdi). The segregating populations,single plant progenies and advancedbreeding lines have been screened usinggrowth room and field screening techniquesat ICRISAT, PAU, Govind Ballab PantUniversity of Agriculture and Technology(GBPUA&T), Pantnagar, and RegionalAgriculture Research Station (RARS),

Figure 14. Field screening technique using perfo-irrigation system for botrytis gray mold development.

14

Table 3. Reaction of wild Cicerspecies to botrytis gray mold (BGM) and ascochyta blight (AB) in greenhousescreening1 at Punjab Agricultural University (PAU), Ludhiana, Punjab, India, 1992–1997.

Disease rating2

Wild Cicer species/accessions Source BGM AB

C. judaicum 182 ICRISAT 3.0 2.0C. judaicum ILWC 19-2 ICARDA 1.5 3.5C. pinnatifidum 188 ICRISAT 1.5 2.0C. pinnatifidum 189 ICRISAT 2.0 2.0C. pinnatifidum 199 ICRISAT 3.0 2.0C. pinnatifidum ILWC 9/S-1 ICARDA 2.0 2.0C. bijugum ILWC 7/S-1 ICARDA 2.0 3.0C. echinospermum ILWC 35/S-1 ICARDA 1.0 1.0C. echinospermum ILWC 39 ICARDA 2.0 3.0

1. Growth room.2. BGM and AB were scored on 1 9 rating scale, where 1 = no disease and 9 = ≥75% plants killed.

Ishurdi, Bangladesh. More than 12 000chickpea germplasm and advanced breedinglines have been screened with little successbecause high level of resistance is notavailable in cultivated genotypes of chickpea(C. arietinum).

Wide hybridization

Cicer judaicum 182 and C. pinnatifidum188 had higher levels of resistance to BGMthan cultivated chickpea during the epidemicyears of 1980 82 (Singh et al. 1982). Later,16 accessions of eight wild Cicer specieswere also screened for resistance to BGMusing growth room technique. Accessionsof C. judaicum, C. bijugum, and C.echinospermum and C. pinnatifidum showedhigh level of resistance to BGM. Incidentally,all these lines also had higher degree ofresistance to ascochyta blight (AB) (Table 3).

At PAU and International Center forAgricultural Research in Dryland Agriculture(ICARDA), crosses were made with someof these wild Cicer species, such as C.pinnatifidum, C. judaicum, and C. bijugum.The breeding materials derived from thesecrosses are in various stages of testing from

F2 to F5, BC1, BC2, and BC3. Chickpea linesGL 92162, GL 92165, and GLK 96239 arebeing tested in yield trials. Several lines arein preliminary trials at PAU. A list of otherchickpea lines derived from the widehybridization and their resistant parents aregiven in Table 4. A total of 837 progenies werederived from the wide hybridization crossesof BG 256 C. judaicum 182, BG 256 C.echinospermum 204, L 550 C. pinnatifidum188, BG 256 C. pinnatifidum 188, andGLK 88012 C. pinnatifidum 188. Theseprogenies were further tested for BGM andAB.

Intraspecific hybridization

From 1980 to 1997, a total of 10 506chickpea germplasm and advanced breedinglines were screened in the growth room forresistance to BGM. During this period,several lines having a BGM rating rangingfrom 3.0 to 5.0 were identified andextensively used in the breeding program.Advanced breeding lines were tested forresistance to BGM at PAU, whereasbreeding material in various segregatinggenerations was screened under field

15

Table 5. Disease scores of the lines most resistant to BGM from 1715 chickpea lines tested in a growthroom at Punjab Agricultural University (PAU), Ludhiana, 1994–97.

BGM score1 No. of lines Chickpea lines

4.0 8 GL 90178, GL 92162, GL 92165, FG 575,FG 576, GN G737, BDG 74, GICG 92228

4.5 4 ICCX 860007-14PN-BPN-BPN-BPN,ICCX 860009-B-BH-2PN-BPN-BPN-BPN,ICCX 860732-BP-BH-BH-BH-1H-2BH,ICCX 870207-BP-BH-4H-BH

5.0 24 GL 84041, GL 90236, GL 90270, GL 94022,GL 94046, GL 94087, GL 94088, GL 94098,GLK 96239, FG 559, FG 579, FG 580, FG 581,FG 596, FG 601, FG 729, SAKI 93-130,PGK 502, ICCX 800837, ICCX 860418-BP-BH-5H-BH, ICCX 860444-BP-5PN-BPN-2H-BH,ICCX 860510-BP-2H-BP, ICCX 880354-BU-BP-51H-BP-BH, ICCX 880355-BH-BP-5H-BH

BGM score on 1 9 scale, where 1 = no disease and 9 = ≥ 75% plants killed.

Table 4. Botrytis gray mold (BGM) scores of chickpea accessions (including wild Cicer spp) and linesderived through wide hydridization and intraspecific crosses at Punjab Agricultural University (PAU),Ludhiana, 1980–831.

Resistant parent BGM rating2 Entry BGM rating

Wide hybridizationC. pinnatifidum 188 1.5 GL 92162 4.0C. echinospermum 204 6.0 GL 92165 4.0C. judaicum 182 3.0 GLK 96239 5.0

Intraspecific hybridizationNegro 4.0 GL 84060 5.0E 100Y 3.0 GL 84107 4.0E 100Y 3.0 GL 84108 4.0P 1528-1-1 4.0 GL 84133 3.0P 1528-1-1 4.0 GL 84295 4.5C 8 4.0 GL 88358 4.5E l00Y 3.0 GL 91040 3.0P 1528-1-1 4.0 GL 85103 4.0

1. Under field conditions in epidemic years 1980 83.2. BGM was scored on 1 9 scale, where 1 = no disease, and 9 = ≥ 75% plants killed.

16

conditions at GBPUA&T, Pantnagar, andRARS, Ishurdi, Bangladesh (Bakr et al. 1997).During 1994/95 and 1996/97, 1715germplasm and advanced breeding linesfrom breeding trials such as Single SelectionTrial (SST), Advanced Yield Trial (AYT),and Field Yield Trial (FYT) were tested forresistance to BGM under controlledconditions in the growth room at PAU.Similarly, other research centers in Indiaincluding GBPUA&T and ICRISAT havealso tested chickpea lines for resistance toBGM (Rathi et al. 1984). Of the 1715 linesscreened, 16 lines were found to be resistant(4.0 4.5 rating), while 22 lines wereidentified as moderately resistant (~5 rating)to BGM (Table 5). In Bangladesh, ICCX860732-BP-BH-BH-1H-BH had a rating of4.0 and ICCX 860444-BP-5PN-BPN-2H-BH a rating of 5.0 for BGM (Bakr et al.1997).

Management by chemicals

Botrytis cinerea, the causal organism ofBGM of chickpea, is internally seedborne.The externally seedborne nature of thefungus is also evident (Grewal et al. 1992,Laha and Grewal 1983b). The seedbornenature of this fungus on chickpea has beenreported by several workers (Cother 1977,Laha and Grewal 1983b, Haware et al.1986, Sandhu and Sah 1988). Themanagement of BGM with chemicalsinvolves seed treatment or foliar sprays withone or a combination of two of the severalfungicides identified to control the disease.

Seed treatment

Seed treatment with fungicide protects thecrop from seedborne inoculum whichcould be the primary source of BGM

development. Seed treatment with amixture of 25% Bavistin (carbendazim) and50% thiram @ 0.25% eliminated externaland internal inoculum of B. cinerea (Grewal1982, Laha and Grewal 1983b). Grewal andLaha (1983) also observed that seedtreatment with carbendazim + thiram, orvinclozolin (Ronilan ), or carbendazimalone @ 0.2% (2 g fungicide kg-1 seed)completely eradicated B. cinerea in chickpeaseeds. Madhu Meeta et al. (1986b) alsostudied the efficacy of different fungicidesfor seed treatment to control BGM andfound that Bavistin was the most effective.Singh and Bhan (1986b) reported that 95%of the seedborne inoculum was eradicatedby seed treatment with triadimefon @0.1%. Singh and Kaur (1990) confirmed theefficacy of triadimefon @ 0.1% for seedtreatment to control BGM. Grewal et al.(1992) observed that soaking of infectedseed for 3 h in a 0.1% suspension of amixture of 25% carbendazim and 50%thiram can reduce seedborne inoculum.These workers also observed that soakingseeds in this fungicide mixture also protectedagainst aerial infection of BGM for about 8weeks after emergence. Bakr et al. (1993)reported that seed treatment alone wasineffective in controlling BGM. Singh et al.(1997) reported that seed treatment withBavistin + thiram (1:1), Indofil M-45,thiabendazole, Ronilan , Rovral , Bavistin@ 0.3% controls seedborne inoculum of B.cinerea. Heat treatment of freshly harvestedseed at 50 C for 5 min, or storage of theseed at 20 C for 12 months reduced thefrequency of occurrence of B. cinerea(Burgess et al. 1997b)

Foliar sprayBotrytis cinerea attacks all aerial parts ofchickpea. Flowers and young fruits are most

17

vulnerable to this disease. Fungicides appliedat regular intervals from the appearance of thefirst symptom can provide considerablecontrol of the disease(Fig. 15). Whenconditions are favorable for BGMdevelopment, effective disease control can beachieved with seed treatment followed by foliarspray with vinclozolin (0.1%) or carbendazim+ thiram (1:1, 0.1%), or carbendazim alone(0.2%) when the first symptoms appear(Grewal and Laha 1983). Madhu Meeta et al.(1986a) observed that four fungicidal sprayswith Bavistin along with seed treatment withthe same chemical reduced BGM incidenceand resulted in higher yields. Rewal andGrewal (1989b) reported that Bavistin 50 WPor Bavistin + thiram gave very goodprotection from aerial infection even whensprayed at 14-day interval.

Pandey (1988) observed that seedtreatment with Bavistin + thiram (1:4)followed by three sprays of Bavistin at 10-day interval provided excellent disease

control that resulted in higher grain yields.Singh and Kaur (1990) reported that seedtreatment with triadimefon (0.1%),carbendazim + thiram (0.3%), Dithane M-45 (0.3%) or Baytan (0.1%), together withone foliar spray of Dithane M-45 ,Hexacap , thiram, thiabendazole, Baytan ,or triadimefon at 50 days after sowing or atthe appearance of first symptomscompletely controlled both primary andsecondary infection of BGM. Two foliarsprays of vinclozolin (0.2%) at flowering andpodding stages along with other agronomicaland cultural management practices discussedabove, reduce BGM incidence and increasegrain yields (Reddy et al. 1990, Haware andMcDonald 1992).

Bakr et al. (1993) observed that seedtreatment (carbendazim 25% and thiram50%, 1:1) together with two foliar sprayswith carbendazim at 14-day interval gavegood protection against BGM in Bangladesh.Haware and McDonald (1993) suggested

Figure 15. Management of botrytis gray mold using fungicide application.

Fungicide sprayed

Fungicide not sprayed

Fungicide sprayed

Fungicide not sprayed

18

the judicious use of vinclozolin (0.2%) in theintegrated management of BGM. Singh etal. (1997) reported that foliar spray withIndofil M-45 , thiabendazole, Baytan ,Bayleton , or thiram during crop growthstages in Feb Mar controlled foliar infection.Haware et al. (1997), and Shrestha et al.(1997) reported that one spray withvinclozolin (0.2%) at the time of flowering inthe integrated management system reducedBGM incidence. However, Bakr et al. (1997)observed that two foliar sprays withvinclozolin (0.2%) was required to controlBGM in Bangladesh. Generally, one spray atflowering followed by another spray after 10days on a moderately resistant chickpeacultivar provided the best results in terms ofdisease management and high grain yield.

Management using biologicalagents

Although repeated fungicide application canalone achieve effective management of BGMin chickpea, biological control of B. cinereausing species of Trichoderma has beenreported in some fruit and vegetable crops(Tronsmo 1986, Nelson and Powelson 1988,

Elad 1994). D’Ercole et al. (1988) obtainedsatisfactory results in controlling B. cinereaof strawberry with Trichoderma species. Verylimited work has been done on biologicalcontrol of B. cinerea using species ofTrichoderma in chickpea. Integrating abiocontrol agent with sublethal doses offungicide seems to be very promising incontrolling plant pathogens withoutdisturbing the biological equilibrium.

Mukherjee and Haware (1993) isolatedspecies of Trichoderma from the rhizosphereof chickpea, tested them against B. cinereain the laboratory, and identified the mosteffective isolate of Trichoderma viride incontrolling BGM of chickpea (Fig. 16).Mukherjee et al. (1995) in further tests,isolated fungicide- (vinclozolin) tolerantisolates for use along with vinclozolin inintegrated BGM management system.Haware et al. (1997) reported that therewere no differences in BGM incidence andgrain yield of chickpea between three spraysof T. viride (107 108 spores mL-1) and threesprays of vinclozolin (0.2%). These Resultsare presented in Table 6. Burgess et al.(1997b) reported that seed treatment withGliocladium roseum suppressed thesporulation of B. cinerea on chickpea seed.

Figure 16. Biological control of botrytis gray mold using Trichoderama viride under greenhouse environment.

19

In the third working group meeting todiscuss collaborative research on botrytisgray mold of chickpea 15 17 April 1996, atPantnagar, Uttar Pradesh, India, it wasunanimously agreed to use the biologicalcontrol agent, T. viride in integratedmanagement of BGM of chickpea.

Integrated ControlChickpea BGM lends itself to integratedcontrol that involves the use of two or moremethods to bring about a reduction indisease incidence (Pande et al. 1998). Agood knowledge of disease epidemiology andcontrol options available to farmers in agiven area are needed before an integratedpackage can be implemented (Fig. 17).Integrated management of the diseaseinvolves chemical control (e.g., seedtreatment with Bavistin + thiram and foliarspray with Bavistin ) and host-plantresistance. The combination of controlmethods can be mutually beneficial. For

example, Pande et al. (1998) suggested thatthe use of seed treatment and need-basedfoliar spray with fungicides could extend thelife of host resistance in the chickpea cultivarAvarodhi and result in low diseasedevelopment. The final methods chosenmust depend on their effectiveness in aparticular situation, and on the farmers’ability to use them.

ConclusionAlthough chickpea research workers havetried to generate adequate information onBGM of chickpea, efficient and completecontrol measures are not yet available foruse by resource-poor farmers in South Asiaand elsewhere. Substantial progress has beenmade in the development of screeningtechniques to identify resistant sources, butthere has been little success, especially inidentifying high levels of resistance incultivated Cicer species. However, high andstable levels of resistance have been

Table 6. Effect of Trichoderma viride on severity of botrytis gray mold (BGM), and yield in BGMnursery, G.B. Pant University of Agriculture and Technology, Pantnagar, India, 1994/95.

Disease rating YieldTreatment (1 9 scale)1 (t ha-1)

Control (no spray) 5.8 1.2

Three sprays of Trichoderma (107 108 spores mL-1) 4.0 1.7at 20-day interval

Three sprays of Ronilan (0.2%) at 20-day interval 4.0 1.8

First spray with Trichoderma (107 108 spore mL-1) + 4.6 1.6Ronilan 0.1%; second spray with Trichoderma(107 108 spore mL-1); third spray with Trichoderma(107 108 spore mL-1) + Ronilan 0.1%

SE ±0.20 ±0.08 CV (%) 9.1 10.1

1. BGM score on a 1 9 rating scale, where 1 = no disease and 9 = ≥75% plants killed.

Source: Haware et al. (1997).

Non-IDM

20

identified in the wild species. It is expectedthat BGM resistant and high-yieldingcultivars endowed with other desirablecharacteristics will be made available tofarmers in the near future. Meanwhile, highpriority should be given to participatory on-farm validation of the available components(including moderate levels of host-plantresistance with high yield) of diseasemanagement and their integration.Extension, and supplying the seed ofavailable resistant cultivars to farmersshould be mandatory for any research anddevelopment program. It is expected thatcollaboration in research on BGM ofchickpea through the Botrytis Gray MoldWorking Group, will increase among themember countries and that thiscollaboration will provide solutions tolocation or region specific problems andcontinue to improve BGM control.

Future Prospects – Use ofTransgenicsTransgenic crops are a hope for fightingdiseases for which genetic resistance eitherdoes not occur or has not been identified.Genetic resistance to B. cinerea in chickpeafalls in this category. Research conducted atScottish Crops Research Institute (SCRI),UK has shown that immature fruits ofraspberry contain a protein, polygalac-turonase-inhibiting protein (PGIP). Thisprotein is highly effective against endo-polygalacturonases enzymes that play amajor role in the invasion of chickpea plantsby B. cinerea. The gene encoding PGIP inraspberry (rPGIP) and also in Kiwifruit(kPGIP) have been isolated. In acollaborative project between SCRI andICRISAT, funded by DFID, these genes havebeen transferred into two chickpea cultivars

Figure 17. Comparison of integrated (IDM) and non-integrated (non-IDM) botrytis gray mold management.

IDM

Non-IDM

21

H-208 and ICCV 5 (SCRI, 2000). Thetransgenic seeds of these two cultivars willnow be evaluated under controlledconditions and later in the field for theirefficacy to control BGM. However, even ifthese genes provide a low level of diseasecontrol, but higher than the available geneticresistance, PGIP-transgenic chickpea canstill be used in an integrated diseasemanagement program.

ReferencesAhmad, Q. 1989. Field screening ofchickpea varieties and cultures against graymold (Botrytis cinerea) in Bihar, India.International Chickpea Newsletter 21:23.Bakr, M.A., and Ahmed, F. 1992. Botrytis graymold of chickpea in Bangladesh. Pages 10 12in Botrytis gray mold of chickpea (Haware,M.P., Faris, D.G., and Gowda, C.L.L., eds.).Patancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics.

Bakr, M.A., Hossain, M.S., and Ahmed, A.U.1997. Research on botrytis gray mold ofchickpea in Bangladesh. Pages 15 18 in Recentadvances in research on botrytis gray mold ofchickpea (Haware, M.P., Lenné, J.M., andGowda, C.L.L., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Bakr, M.A., Rehman, M.M., Ahmed, F., andKumar, J. 1993. Progress in the managementof botrytis gray mold of chickpea in Bangladesh.Pages 17 19 in Recent advances in researchon botrytis gray mold of chickpea (Haware,M.P., Gowda, C.L.L., and McDonald, D., eds.).Patancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics.

Bakr, M.A., Rehman, M.H., Hossain, M.S.,and Ahmed, A.U. 1998. Steps towardsmanagement of botrytis gray mold of chickpeain Bangladesh. Pages 15 18 in Recent advancesin research and management of botrytis graymold of chickpea: summary proceedings of theFourth Working Group Meeting to DiscussCollaborative Research on Botrytis Gray Moldof Chickpea, 23 26 Feb 1998, BARI,Joydebpur, Gazipur 1701, Bangladesh (Pande,S., Bakr, M.A., and Johansen, C., eds.).Patancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics.

Bhan, U., and Chatrath, M.S. 1994. Acquiredresistance to carbendazim in Botrytis cinereaPers. Ex. Fr. Crop Research 8(1):162 168.

Brinsmead, R.B. 1992. Chickpea cultivar ×planting time studies in Queensland, Australia.Page 64 in Program and abstracts, SecondInternational Food Legume ResearchConference, 12 16 Apr 1992, Cairo, Egypt.Cairo, Egypt: International Food LegumesResearch Conference Organizing Committee.

Burgess, D.R., Bretag, T., and Keane, P.J.1997a. Seed to seedling transmission ofBotrytis cinerea in chickpea and disinfestationof seed with moist heat. Australian Journal ofExperimental Agriculture 37(2):223 229.

Burgess, D.R., Bretag, T., and Keane, P.J.1997b. Biocontrol of seedborne Botrytiscinerea in chickpea with Gliocladium roseum.Plant Pathology 46(3):298 305.

Butler, D.R. 1993. How important is cropmicroclimate in chickpea botrytis gray mold?Pages 7 9 in Recent advances in research onbotrytis gray mold of chickpea (Haware, M.P.,Gowda, C.L.L., and McDonald, D., eds.).Patancheru 502 324, Andhra Pradesh, India:

22

International Crops Research Institute for theSemi-Arid Tropics.

Carranza, J.M. 1965. [Wilt of chickpea (Cicerarietinum L.) caused by B. cinerea] (In Es.).Revista de la Facultad de Agronomia,Universidad Nacional de la Plata. 41:135 138.

Coley-Smith, J.R. 1980. Sclerotia and otherstructures in survival. Pages 85 114 in TheBiology of Botrytis (Coley-Smith, J.R.,Verhoeff, K., and Jarview, W.R., eds.). London:Academic Press.

Cother, E.J. 1977. Isolation of importantpathogenic fungi from seeds of chickpea. SeedScience and Technology 5:593.

D’Ercole, N., Nipoti, P., Finessi, and L.E.,Manzali, D. 1988. Review of several years ofresearch in Italy on the biological control ofsoil fungi with Trichoderma spp. OrganizationEuropeenne et Mediterraneenne pour laprotection des plants Bulletin 18(1):95 102.

Elad, Y. 1994. Biological control of grape graymold by Trichoderma harzianum. CropProtection 13:35 38.

Ellis, M.B., and Waller, J.M. 1974. Sclerotiniafuckeliana. CMI Descriptions of PathogenicFungi and Bacteria No. 431. Kew, UK:Commonwealth Mycological Institute.

Grewal, J.S. 1982. Control of importantseedborne pathogens of chickpea. IndianJournal of Genetics and Plant Breeding42:393 398.

Grewal, J.S. 1988. Diseases of pulse crops: anoverview. Indian Phytopathology 41(1):1 14.

Grewal, J.S., and Laha, S.K. 1983. Chemicalcontrol of botrytis blight of chickpea. IndianPhytopathology 36:516 520.

Grewal, J.S., Pal, M., and Rewal, N. 1992.Botrytis gray mold of chickpea in India. Pages

6 9 in Botrytis gray mold of chickpea (Haware,M.P., Faris, D.G., and Gowda, C.L.L., eds.).Patancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics.

Grooves, J.W., and Loveland, C.A. 1953. Theconnection between Botryotinia fuckeliana andBotrytis cinerea. Mycologia 45:415 425.

Haware, M.P., and McDonald, D. 1992.Integrated management of botrytis gray moldof chickpea. Pages 3 6 in Botrytis gray mold ofchickpea (Haware, M.P., Faris, D.G., andGowda, C.L.L., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Haware, M.P., and McDonald, D. 1993. Botrytisgray mold of chickpea. Pages 3 6 in Recentadvances in research on botrytis gray mold ofchickpea (Haware, M.P., Gowda, C.L.L., andMcDonald, D., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Haware, M.P., and Nene, Y.L. 1982. Screeningchickpea for resistance to botrytis gray mold.International Chickpea Newsletter 6:17 18.

Haware, M.P., Narayana Rao, J., and Pundir,R.P.S. 1992. Evaluation of wild Cicer speciesfor resistance to four chickpea diseases.International Chickpea Newsletter 27:16 18.

Haware, M.P., Nene, Y.L., and Mathur, S.B.1986. Seedborne diseases of chickpea.Technical Bulletin no. 1. Copenhagen,Denmark: Danish Government Institute ofSeed Pathology for Developing Countries, andPatancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics. 32 pp.

Haware, M.P., Tripathi, H.S., Rathi, Y.P.S.,Lenn , J.M., and Jayanthi, S. 1997. Integrated

23

management of botrytis gray mold of chickpea:cultural, chemical, biological, and resistanceoptions. Pages 9 12 in Recent advances inresearch on botrytis gray mold of chickpea(Haware, M.P., Lenné, J.M., and Gowda,C.L.L., eds.). Patancheru 502 324, AndhraPradesh, India: International Crops ResearchInstitute for the Semi-Arid Tropics.

Joshi, M.M., and Singh, R.S. 1969. A botrytisgray mold of gram. Indian Phytopathology22:125 128.

Joshi, S. 1992. Botrytis gray mold of chickpeain Nepal. Pages 12 13 in Botrytis gray mold ofchickpea (Haware, M.P., Faris, D.G., andGowda, C.L.L., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Karki, P.B., Joshi, S., Chaudhary, G., andChaudhary, R.N. 1993. Studies on botrytis graymold of chickpea in Nepal. Pages 11 13 inRecent advances in research on botrytis gray moldof chickpea (Haware, M.P., Gowda, C.L.L., andMcDonald, D., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Karki, P.B., Tiwari, K.R., Onkar Singh, andBharati, M.P. 1989. Effect of sowing date onthe incidence of botrytis gray mold in chickpea.International Chickpea Newsletter 21:21 23.

Kharbanda, P.D., and Bernier, C.C. 1979.Botrytis gray mold of chickpea in Manitoba.Plant Disease Reporter 63:662.

Laha, S.K., and Grewal, J.S. 1983a. A newtechnique to induce abundant sporulation inBotrytis cinerea. Indian Phytopathology36:409.

Laha, S.K., and Grewal, J.S. 1983b. Botrytisblight of chickpea and its perpetuation throughseed. Indian Phytopathology 36:630 634.

Laha, S.K., and Khatua, D.C. 1988. Sourcesof resistance against gray mold andstemphylium blight of gram. Environment andEcology 6(3):656 659.

Madhu Meeta, Bedi, P.S., and Jindal, K.K.1986a. Survival of Botrytis cinerea causalorganism of gray mold of gram in Punjab. IndianJournal of Plant Pathology 4(2):143 145.

Madhu Meeta, Bedi, P.S., and Jindal, K.K.1988. Host range of Botrytis cinerea theincitant of grey mould of gram. Plant DiseaseResearch 3(1):77 78.

Madhu Meeta, Bedi, P.S., and Krishna Kumar.1986b. Chemical control of gray mold of gramcaused by Botrytis cinerea in Punjab. Journalof Research, Punjab Agricultural University23(3):435 438.

Mahmood, M., and Sinha, B.K. 1990. Graymold disease of bengal gram in Bihar. Finaltechnical bulletin. Dholi, Bihar, India: TirhutCollege of Agriculture, 35 pp.

Malik, B.A., Iqbal, S.M., and Bashir, M. 1993.Present status and future research outlook forbotrytis gray mold a potential threat tochickpea in Pakistan. Pages 19 20 in Recentadvances in research on botrytis gray mold ofchickpea: summary proceedings of the SecondWorking Group Meeting to DiscussCollaborative Research on Botrytis Gray Moldof Chickpea, 14 17 Mar 1993, Rampur,Nepal. Patancheru 502 324, Andhra Pradesh,India: International Crops Research Institutefor the Semi-Arid Tropics.

Micheli, P.A. 1929. Nova Platarum Generajuxta Tournefortii methodum disposite.Florence, 234 pp.

Mukherjee, P.K., and Haware, M.P. 1993.Biological control of botrytis gray mold ofchickpea. International Chickpea Newsletter28:14 15.

24

Mukherjee, P.K., Haware, M.P., and Jayanthi,S. 1995. Preliminary investigations onintegrated biocontrol of botrytis gray mold ofchickpea. Indian Phytopathology48(2):141 149.

Nelson, M.E., and Powelson, M.L. 1988.Biological control of gray mold of snap bean byTrichoderma hamatum. Plant Disease72:727 729.

Nene, Y.L., Sheila,V.K, and Sharma, S.B.1984. A world list of chickpea (Cicerarietinum) and pigeonpea [Cajanus cajan (L.)Millsp.] pathogens. ICRISAT Pulse PathologyProgress Report 32. Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics(Limited distribution).

Pande, S. 1998. Diseases of chickpea in Nepaland Bangladesh A Survey Report. Trip ReportJan 1998. Patancheru 502 324, AndhraPradesh, India: International Crops ResearchInstitute for the Semi-Arid Tropics. 3 pp(Limited circulation).

Pande, S., Johansen, C., and Narayana Rao,J. 1998. Management of botrytis gray mold ofchickpea a review. Pages 23 40 in Recentadvances in research and management ofbotrytis gray mold of chickpea (Pande, S., Bakr,M.A., and Johansen, C., eds.). Patancheru 502324, Andhra Pradesh, India: InternationalCrops Research Institute for the Semi-AridTropics. 52 pp.

Pande, S., Sharma, S.B., and Ramakrishna, A.2000. Biotic stresses affecting legumesproduction in the Indo-Gangetic Plain. Pages129 155 in Legumes in rice and wheat croppingsystems of the Indo-Gangetic Plain Constraintsand Opportunities (Johansen, C., Duxbury,J.M., Virmani, S.M., Gowda, C.L.L., Pande,S., and Joshi, P.K., eds.). Patancheru 502 324,

Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics,and Ithaca, New York, USA: Cornell University.

Pandey, B.K. 1988. Studies on botrytis graymold of chickpea (Cicer arietinum L.). Ph.D.thesis, Govind Ballabh Pant University ofAgriculture and Technology, Pantnagar, UttarPradesh, India.

Rathi, Y.P.S., and Tripathi, H.S. 1991. Hostrange of Botrytis cinerea Pers. ex Fr., the causalagent of gray mold of chickpea. InternationalChickpea Newsletter 24:37 38.

Rathi, Y.P.S., Tripathi, H.S., Chaube, H.S.,Beniwal, S.P.S., and Nene, Y.L. 1984.Screening chickpea for resistance to botrytisgray mold. International Chickpea Newsletter11:31 33.

Reddy, M.V., Ghanekar, A.M., Nene, Y.L.,Haware, M.P., Tripathi, H.S., and Rathi, Y.P.S.1993. Effect of vinclozolin spray, plant growthhabit and interrow spacing on botrytis gray moldand yield of chickpea. Indian Journal of PlantProtection 21(1):112 113.

Reddy, M.V., Nene, Y.L., Singh, G., andBashir, M. 1990. Strategies for managementof foliar diseases of chickpea. Pages 117 127in Chickpea in the nineties: proceedings of theSecond International Workshop on ChickpeaImprovement 4-8 Dec 1989, ICRISAT, India.(van Rheenen, H.A., Saxena, M.C., Walley,B.J., and Hall, S.D., eds.). Patancheru 502324, Andhra Pradesh, India: InternationalCrops Research Institute for the Semi-AridTropics.

Reddy, M.V., Onkar Singh, Bharati, M.P., Sah,R.P., and Joshi, S. 1988. Botrytis gray moldepiphytotic of chickpea in Nepal. InternationalChickpea Newsletter 19:15.

Rewal, N., and Grewal, J.S. 1989a. Effect oftemperature, light and relative humidity on

25

conidial germination of three strains of Botrytiscinerea infecting chickpea. IndianPhytopathology 42(1):79 83.

Rewal, N., and Grewal, J.S. 1989b.Differential response of chickpea to greymould. Indian Phytopathology 42(2):265 268.

Sandhu, R., and Sah, D.N. 1988. Effect ofbotrytis gray mold on chickpea flowers, podformation and yield in Nepal. InternationalChickpea Newsletter 10:13.

SCRI. (Scottish Crops Research Institute).2000. Enhancement of resistance to Botrytisgray mold of chickpea using pGIP genes.Holdback project R6409(H). Final Report. 1Oct 1996 to 31 Jan 2000.

Shaw, F.J.F. and Ajrekar, S.L. 1915. The genusRhizoctonia in India. Memoirs of theDepartment of Agriculture in India BotanicalSeries 7:177.

Shrestha, K., Joshi, S., Chaudhary, G.,Chaurasia, P.C., and Adhikary, R.C. 1997.Research on botrytis gray mold of chickpea inNepal. Pages 19 20 in Recent advances inresearch on botrytis gray mold of chickpea(Haware, M.P., Lenné, J.M., and Gowda,C.L.L., eds.). Patancheru 502 324, AndhraPradesh, India: International Crops ResearchInstitute for the Semi-Arid Tropics.

Singh, G. 1997. Epidemiology of botrytis graymold of chickpea. Pages 47 50 in: Recentadvances in research on botrytis gray mold ofchickpea (Haware, M.P., Lenné, J.M., andGowda, C.L.L., eds.). Patancheru 502 324,Andhra Pradesh, India: International CropsResearch Institute for the Semi-Arid Tropics.

Singh, G., and Bhan, L.K. 1986a. Physiologicalraces of Botrytis cinerea causing gray mold ofchickpea. Plant Disease Research 1:69 74.

Singh, G., and Bhan, L.K. 1986b. Chemicalcontrol of gray mold in chickpea. InternationalChickpea Newsletter 15:18 20.

Singh, G., and Kapoor, S. 1984. Role ofincubation and photoperiod on the intensityof botrytis gray mold of chickpea. InternationalChickpea Newsletter 12:23 24.

Singh, G., Kapoor, S., and Singh, K. 1982.Screening of chickpea for gray mold resistance.International Chickpea Newsletter 7:13 14.

Singh, G., and Kaur, L. 1989. Geneticvariability studies and scope for improvementin chickpea, Punjab, India. InternationalChickpea Newsletter 20:7.

Singh, G., and Kaur, L. 1990. Chemicalcontrol of gray mold of chickpea. Plant DiseaseResearch 5(2):132 137.

Singh, G., Kaur, L., and Sharma, Y.R. 1991.Ascochyta blight and gray mold resistance inwild species of Cicer. Crop Improvement18(2):150-151.

Singh, G., Kaur, L., and Sharma, Y.R. 1992.Exploitation of host-plant resistance to managebiotic stresses in chickpea. Page 76 in Programand abstracts, Second International FoodLegume Research Conference, 12 16 Apr1992. Cairo, Egypt.

Singh, G., Kumar, B., and Sharma, Y.R. 1997.Botrytis gray mold of chickpea in Punjab, India.Pages 13 14 in Recent advances in researchon botrytis gray mold of chickpea (Haware,M.P., Lenné, J.M., and Gowda, C.L.L., eds.).Patancheru 502 324, Andhra Pradesh, India:International Crops Research Institute for theSemi-Arid Tropics.

Singh, G., Sharma, Y.R., and Bains, T.S. 1998.Status of botrytis gray mold of chickpea researchin Punjab, India. Pages 7 14 in Recent

26

advances in research and management ofbotrytis gray mold of chickpea: Summaryproceedings of Fourth Working Group Meetingto Discuss Collaborative Research on BotrytisGray Mold of Chickpea, 23 26 Feb 1998,BARI, Joydebpur, Gazipur 1701, Bangladesh(Pande, S., Bakr, M.A., and Johansen, C.,eds.). Patancheru 502 324, Andhra Pradesh,India: International Crops Research Institutefor the Semi-Arid Tropics.

Singh, M.P. 1989. Studies on survivability ofBotrytis cinerea Pers. ex Fr, causal agent of greymould of chickpea. M.S. thesis, Govind BallabPant University of Agriculture and Technology,Pantnagar, Uttar Pradesh, India.

Singh, M.P., and Tripathi, H.S. 1992. Effectof temperature and depth of burial on survivalof Botrytis cinerea Pers. ex Fr., causal organismof grey mould of chickpea. Indian Journal ofMycology and Plant Pathology 22(1):39 43.

Singh, M.P., and Tripathi, H.S. 1993. Effectof storage temperatures on the survival ofBotrytis cinerea in chickpea seeds. IndianJournal of Mycology and Plant Pathology23(2):177 179.

Singh, R.B. 1970. Studies on the control of blightand wilt of gram. M.Sc. thesis. Govind BallabPant University of Agriculture and Technology,Pantnagar, Uttar Pradesh, India. 49 pp.

Tripathi, H.S., and Rathi, Y.P.S. 1992.Epidemiology of botrytis gray mold ofchickpea. Pages 8 9 in Botrytis gray mold ofchickpea. (Haware, M.P., Faris, D.G., Gowda,C.L.L., eds.). Patancheru 502 324, AndhraPradesh, India: International Crops ResearchInstitute for the Semi-Arid Tropics.

Tronsmo, A. 1986. Trichoderma used as abiocontrol agent against Botrytis cinerea rots onstrawberry and apple. Scientific Reports fromthe Agricultural University of Norway 65:1 22.

About ICRISAAbout ICRISAAbout ICRISAAbout ICRISAAbout ICRISATTTTT

The semi-arid tropics (SAT) encompasses parts of 48 developing countries including most ofIndia, parts of southeast Asia, a swathe across sub-Saharan Africa, much of southern and easternAfrica, and parts of Latin America. Many of these countries are among the poorest in the world.Approximately one-sixth of the world’s population lives in the SAT, which is typified byunpredictable weather, limited and erratic rainfall, and nutrient-poor soils.

ICRISAT’s mandate crops are sorghum, pearl millet, finger millet, chickpea, pigeonpea, andgroundnut; these six crops are vital to life for the ever-increasing populations of the semi-aridtropics. ICRISAT’s mission is to conduct research which can lead to enhanced sustainableproduction of these crops and to improved management of the limited natural resources of theSAT. ICRISAT communicates information on technologies as they are developed throughworkshops, networks, training, library services, and publishing.

ICRISAT was established in 1972. It is one of 16 nonprofit, research and training centers fundedthrough the Consultative Group on International Agricultural Research (CGIAR). The CGIARis an informal association of approximately 50 public and private sector donors; it is co-sponsored by the Food and Agriculture Organization of the United Nations (FAO), the UnitedNations Development Programme (UNDP), the United Nations Environment Programme(UNEP), and the World Bank.