Breeding for drought resistance in chickpea

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Breeding for drought resistance in chickpea

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CONTENTS Introduction Mechanism of Drought resistance Screening studies Genetic studies on Drought resistance Genetic variability Source of resistance Breeding approaches Biotechnological approaches Conclusions Future thrust

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INTRODUCTION

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4 Chickpea Area (Million ha) Production (Million tones) Productivity (kg/ha) Gujarat 0.24 0.09 808 India 6.93 5.6 808 World 10.33 7.9 767 Table 1: Area, Production and Productivity in the year (2005-06) FAO (2005) 4

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5 Yield gap of rainfed agriculture in the Semi Arid Tropics Yield losses due to water deficits vary depending on time, intensity and duration of deficits, coupled with high irradiance and temperature. Global yield losses due to drought in chickpea have been estimated to be around 3.7 million tonnes. In addition to direct effect of drought on yield, it also reduces the potential beneficial effects of improved crop management practices such as fertilizer application or intercropping in terms of increased production. Drought reduces both carbon assimilation and symbiotic nitrogen fixation. 5

Table 2: Production losses in chickpea due to stresses. :

6 Table 2: Production losses in chickpea due to stresses. Country Mean Prod. (000 t.) Estimated production losses (000 t.) Abiotic stress Biotic stress Drought/heat Cold Salinity AB BGM V Hel. Total Bangladesh 69 27 0 10 0 17 7 14 75 China 210 126 0 17 - - - 42 185 India 4329 2026 269 569 318 161 501 925 4759 Myanmar 110 55 0 11 0 0 0 22 88 Nepal 17 7 3 0 0 5 1 4 20 Pakistan 455 182 68 114 91 46 91 114 706 Middle East 916 532 133 85 218 0 57 145 1170 Africa 267 160 5 40 19 0 22 44 290 America 171 72 23 39 0 0 30 43 207 Europe 79 47 4 5 10 0 3 8 77 Australia 83 42 13 8 4 4 8 21 138 World 6700 3276 518 898 660 233 720 1382 7715 ICRISAT Kumar et al. (1996) AB – Ascochyta blight, BGM- Botrytis gray mold, V- Virus, Hel- Helicoverpa pod borer 6

What is drought?:

7 What is drought? * Drought : Drought refers to the condition of soil moisture deficiency or water scarcity. * Drought resistance: Drought resistance refers to survival of plant under water deficit or scarcity conditions without injury. 7

Main features of drought:

8 Main features of drought Drought is characterized with soil water deficit or low soil moisture. About 36% of the land area constitutes arid and semi arid zones. Arid and semi arid areas are more prone to drought. Drought leads to reduction in both yield and quality of economic product in crop plants. It has adverse effect on plant growth and development. Drought damages chloroplasts and lowers photosynthetic output. There is an increase in proline level in the leaves of plants which are subjected to all stresses. Drought resistance is a genetically controlled physiological property of plant species. 8

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9 Plant responses to drought are very complex as stress itself involves various climatic, edaphic and agronomic factors, frequently complicated by major variation in time of occurrence and duration intensity The general complexity of drought is often aggravated under the conditions of semi arid tropics by erratic and unpredictable rainfall and by occurrence of high temperature, high level of solar radiation and poor soil characteristics Identifying a plant phenotype which can be used in the breeding programme to transfer drought resistance traits in to cultivars with high yielding genetic back ground . 9

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MECHANISM OF DROUGHT RESISTANCE

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11 TRAITS FOR SUSTAINING YIELD UNDER DROUGHT Yield Potential (under moderate stress) Drought adaptation (under severe stress) 11

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12 Constitutive / Stress Responsive Traits Drought Escape Dehydration Avoidance Dehydration (desiccation) Tolerance

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Drought Escape Drought susceptible variety performs well in drought environment simply by avoiding the period of drought Dehydration Avoidance Ability of plant to retain a relatively higher level of hydration under conditions of soil or atmospheric water stress (Maintenance of turgor and volume) Maintenance of water uptake Reduction of water loss Change in tissue characteristics Dehydration Tolerance Significantly lower level of change are induced in it than those in another genotype when both are subjected to the same level of hydration. Protoplasmic tolerance

Table 3 : Drought tolerance traits and their usefulness and ease of screening in breeding programme.:

14 Table 3 : Drought tolerance traits and their usefulness and ease of screening in breeding programme. Traits Usefulness Ease of screening Drought escape Phenological development Very high Easy Drought tolerance Early vigour High Easy Transpiration efficiency High Easy, but costly Stomatal control High Difficult Osmotic adjustment High Difficult Deeper and denser roots High Very difficult Membrane stability Low Easy Wembley Turner (2001) 14

Figure 1:Root characters of different genotypes.:

15 Figure 1:Root characters of different genotypes. 15

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16 Describing drought prone conditions in precise and quantitative terms, as relevant to chickpea and area affected by drought. Generating scientific knowledge and information on chickpea responses to drought. Developing reproducible field and laboratory methods for screening and identifying drought tolerant germplasm. Characterizing drought tolerant germplasm for easily identifiable, shoot and root traits, reflective of internal plant mechanisms and process that confers drought resistance. Incorporating drought tolerant traits in agronomically useful genetic background using conventional breeding methodology. Developing tools for rapid incorporation of traits. Primary requirements for drought tolerance 16

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SCREENING STUDIES

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Laboratory method In this method to identify genotypic difference in germinability in laboratory, osmotic solutions like polyethylene glycol(PEG) was used. The osmotic effect of drought are known to be comparable to true drought effects only under non limiting of water movement where the soil and seed contact is perfect. In field condition, it is difficult to visualize a perfect seed and soil contact. Therefore instead of osmotic solutions, soils brought to different moisture tensions and packed in seed germination trays. Results shows that seedlings failed to emerged in vertisol at soil moisture content below 20%. ICRISAT Saxena et al. (1984)

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19 Field method: The field testing was conducted on deep vertisol at ICRISAT centre The field is uniformly irrigated with overhead system using perforated pipes. The seedling is done at uniform depth of 5 cm on different dates, to obtain contrasting difference in soil moisture contents at time of seedling During the course of experiment, no rainfall is received. Counted number of seeds is sown in subplot. Soil moisture at 0-10 cm soil depth was determined gravimetrically at three places. The percentage of seedling that emerge is computed. 19 Continue..

Figure 2: Screening for drought tolerance chickpea using late spring planting in WANA (West Asia & North Africa ):

20 Figure 2: Screening for drought tolerance chickpea using late spring planting in WANA (West Asia & North Africa ) 20

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21 Description and delineation of iso -drought environments Selections in germplasm for adaptation drought Empirical approach Directed approach Multilocational evaluation to select escapes Identification of Avoid/ Tol ./ Resis.irrespective of adaptation Developing field and laboratory methods to create representive and reproducible drought environments Selections for adaptation to the drought environment, total dry matter and yield Morphological, anatomical,physilogical and biochemical characterization Field and laboratory screenings Yield testing of adapted good agronomoic genotypes Drought Avod/ Tol ./Resis Recommendation for cultivation in the drought environment Initiation of plant improvement program for drought resistance Figure 3: Procedure for identifying drought tolerant chickpea genotypes . Saxena and Johansen.(1984) ICRISAT 21

Table 4: Relationship between relative water content (RWC) and Membrane Injury Index(MII) in four genotypes of chickpea 60 day after sowing :

22 Table 4: Relationship between relative water content (RWC) and Membrane Injury Index(MII) in four genotypes of chickpea 60 day after sowing Genotypes Relative water content (%) Time (hrs.) at 40 0 C Membrane injury index Time (hrs.) at 40 0 C 0 2 4 0 2 4 C- 214 79.6 70.8 66.4 0.145 0.231 0.387 BG- 256 81.2 76.0 61.1 0.104 0.206 0.342 Annigeri 77.5 75.0 57.6 0.203 0.435 0.449 ICC-4958 83.6 63.6 56.8 0.202 0.473 0.519 SEm± 0.81 0.72 0.66 0.006 0.007 0.007 CD at 5% 2.41 2.15 1.82 0.017 0.021 0.022 IARI, New Delhi Deshmukh et al .(2002) 22

Table 5: Classification of chickpea genotypes on the basis of membrane injury index at seed filling phase (125 DAS):

23 Table 5: Classification of chickpea genotypes on the basis of membrane injury index at seed filling phase (125 DAS) Genotypes Membrane injury index Tolerant BG- 276 0.24 IPC-10459 0.28 IPC-92-1 0.29 BG-162 0.30 BG-212 0.30 C-325 0.31 BG-364 0.32 BG-365 0.33 PGD- 84- 16 0.33 C-214 0.34 Medium tolerant T- 315 0.35 BG-256 0.38 H-208 0.38 IPC-92-2 0.39 IPC-94-132 0.39 G- 130 0.39 Susceptible BG-372 0.47 Annegeri 0.48 H-75-35 0.52 Hima 0.55 SEm± 0.019 CD at 5% 0.061 IARI, New Delhi Deshmukh et al .(2002) 23

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24 Figure 4: Membrane injury of genotypes at pod formation stage as influenced by time of planting. IARI, New Delhi Deshmukh et al. (2004) 24

Table 6: Screening of drought resistance for seed yield (kg/ha) of genotypes at four different dates of sowing .:

25 Table 6: Screening of drought resistance for seed yield (kg/ha) of genotypes at four different dates of sowing . Entry Dates of planting 28 Feb 10 March 20 March 30 March mean ILC 72 485 130 11 7 158 ILC 3279 570 365 76 37 262 FLIP 85-142C 339 150 3 6 124 ILC 1929 1439 1276 948 737 1100 ILV 482 1215 1033 661 444 838 ILC 1919 1043 863 524 652 770 FLIP 87-5C 1281 1126 776 704 972 FLIP 87-59C 1435 1420 935 1189 1245 ILC 6104 1611 1176 893 1007 1172 ILC 6118 1507 1263 789 963 1131 Mean 1065 920 587 559 783 SE± (Dates) 42.92 SE± (Entry) 67.35 SE± (Dates across entries) 138.78 C.V. % (Date) 33.58 C.V. % (Entry) 21.07 L.S.D. (0.05) Dates 105.03 L.S.D. (0.05) Entries 132.01 L.S.D. (0.05) Dates across entries 278.48 ICARDA, Syria Singh et al. (1997)

Table 7: Performance of top yielding chickpea cultivars under moisture stress conditions:

26 Table 7: Performance of top yielding chickpea cultivars under moisture stress conditions Sr.No. Genotypes Plant height (cm) Days to flowering Days to maturity Pods per plant 100 seed weight (g) Seeds per pod Biomass (2.4m 2 ) (g) Harvest index (%) Seed yield (kg /ha) % superiority over check 1 BG 365 48.11 87.44 139.78 338.4 17.86 1.29 875.2 36.26 2714 31.6 2 BG 364 50.78 90.00 139.44 244.7 25.06 0.92 721.3 41.45 2501 21.3 3 BG 362 50.78 86.00 139.00 249.0 23.78 1.37 741.1 38.52 23.94 16.1 4 BG 391 51.33 90.22 138.33 213.2 22.97 1.15 661.2 39.53 2189 6.2 5 IPC94-132 52.17 92.00 138.78 276.6 20.39 1.16 691.6 37.03 2184 5.9 6 BG 256 54.78 91.11 140.33 217.1 24.30 1.05 687.2 37.05 2157 5.1 7 ICC 4958 (check) 53.00 83.56 140.56 273.9 30.18 0.91 641.7 37.01 2062 - New Delhi Yadav et al. (2004) 26

Table 8: Performance of highest yielding drought resistant genotypes under rainfed and irrigated conditions.:

27 Table 8: Performance of highest yielding drought resistant genotypes under rainfed and irrigated conditions. Entry name Days to flower (rainfed) Rainfed seed yield Drought rating score Irrigated seed yield (kg/ha) Drought susceptibility index ILC 142 48 1426 3 2212 0.744 ILC 391 56 1166 5 1988 0.981 ILC 588 50 1113 4 2176 0.986 ILC 1306 54 1352 4 1950 0.716 ILC 1799 54 1135 4 2376 1.033 ILC 2216 51 1141 6 1952 0.695 ILC 2516 51 1171 4 1858 0.853 ILC 3550 50 1135 4 2055 0.810 ILC 3764 53 1246 4 2310 0.674 ILC 3832 52 1200 4 2022 0.975 ILC3843 49 1332 4 2532 0.765 ILC 4236 50 1064 4 2325 1.046 Continue..

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28 Singh et al . (1997) ICARDA, Syria Drought rating score 1= free from damage, 9= all plants dried without any seed. Entry name Days to flower (rainfed) Rainfed seed yield (kg/ha) Drought resistance score Irrigated seed yield (kg/ha) Drought susceptibility index FLIP 87-7C 49 1100 4 2220 O.961 FLIP 87-8C 49 1016 4 2368 0.969 FLIP 87-58C 46 1085 4 2197 1.069 FLIP 87-59C 48 1191 4 2245 0.995 FLIP 88-42C 49 1392 4 2075 1.043 FLIP 87- 85C 49 1028 4 1858 0.789 ICC 4958 49 1194 5 1931 0.968 Susceptible controls ILC 72 68 74 9 1445 1.644 ILC 1171 75 161 8 931 1.697 Mean of 81 genotypes - 961.4 - 1854.7 - SE ± - 119.2 - 214.4 - C. V.(%) - 21.5 - 20.0 - LSD at P≤ 0.05 - 330.3 - 594.1 -

Table 9: Effect of PEG 6000 on Germination of chickpea cultivars:

29 Table 9: Effect of PEG 6000 on Germination of chickpea cultivars Cultivar % Germination % Reduction over control Distilled water (Control) PEG PEG -5 bar solution -10 bar solution -5 bar solution -10 bar solution HMS- 25 100.0 98.7 9.3 1.3 90.7 BG-244 93.3 54.7 4.0 41.4 95.7 HMS- 30 97.3 80.0 10.7 17.8 89.0 GF- 734 93.3 69.3 6.7 25.7 92.8 H-78- 84 100.0 98.1 1.3 1.3 98.7 GL- 796 77.3 50.7 1.3 34.5 98.3 RSG- 45 100.0 94.7 0.0 5.3 100.0 H- 76 -3 98.7 77.3 1.3 21.6 98.-6 S. E. ± (interaction) - 5.47 - - 95.6 Mean 95.0 78.0 4.3 18.8 - S. E. ± (solutions) - 1.93 - - - S. E. ± (varieties) - 3.16 - - - Dutt et al.( 1982) Udaipur

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30 The following equality suggested by Roeielle and Hamblin was used for determination of tolerance to drought stress (TDS) of genotypes . Where Y 3 is TDS, Y 3 = Y 2 – Y 1 Y 1 is the seed yield in non stress environment (winter sown) and Y 2 seed yield in stress environment (spring -sown). Mean productivity was calculated by using formula : Y 4 = (Y 1 + Y 2 )/2 The mean productivity (MP) was defined as Y 4 and rate of productivity was attributed as Y 5 . Y 5 = (Y 2 / Y 1 ) Also, drought susceptibility index (DSI) was calculated with following equivalent used for cereals and applied Fischer and Maurer, where D is the rate of means as : DSI = 1- (Y 2 / Y 1 )/D D= mean of all of genotypes in Y 2 / mean of all of genotypes in Y 1 30 Turkey Toker et. al . (1997)

Table 10: Varietal response for seed yield (kg/ha), mean productivity, tolerance to drought stress, rate of productivity and drought susceptibility index :

31 Table 10: Varietal response for seed yield (kg/ha), mean productivity, tolerance to drought stress, rate of productivity and drought susceptibility index Lines Yield in Y 1 (kg/ha) Yield in Y 2 (kg/ha) MP TDS RP DSI FLIP 90-8C 1000 964 982 -36 0.964 -1.166 FLIP 90-111C 911 661 786 -250 0.726 -0.632 FLIP 91-15C 1250 1054 1152 -196 0.843 -0.894 FLIP 91- 19C 1107 714 813 -589 0.468 -0.052 FLIP 91- 45C 2161 518 1402 -1518 0.298 0.330 FLIP 91- 50C 1536 643 1295 -482 0.686 -0.542 FLIP 91- 58C 1518 1154 1152 -732 0.518 -0.164 FLIP 91- 59C 1482 786 1232 -500 0.663 -0.490 FLIP 91- 63C 1357 982 1179 -357 0.737 -0.656 FLIP 92- 48C 1000 625 813 -375 0.625 -0.405 FLIP 92- 58C 1232 786 1009 -446 0.638 -0.434 FLIP92- 65C 946 589 768 -357 0.623 0.025 Mean 1340 775 1062 -574 0.590 -0.313 Checks FLIP 82- 150C 1617 1036 1327 -581 0.641 -0.441 ILC 482 1786 446 1116 -1340 0.250 0.438 Urkutlu 1196 750 873 -466 0.627 -0.409 Mean 1533 744 1139 -796 0.504 -0.137 Y 1 =yield under non stress conditions, Y 2 =yield under stress conditions, MP= mean productivity, TDS= tolerance to drought stress, RP= rate of productivity, DSI= drought susceptibility index Turkey Tokar et al. (1996)

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GENETICS OF DROUGHT RESISTANCE

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33 GENETIC VARIABILITY

Figure 5: Effect of water potential on changes in fresh mass of roots during germination.:

34 Figure 5: Effect of water potential on changes in fresh mass of roots during germination. 34 Fresh mass of root (mg) Gupta et al .(1991) Ludhiana control stress control stress : 0 MPa Control, : -0.14MPa stress

Figure 6: Effect of water potential on changes in fresh mass of shoots during germination.:

35 Figure 6: Effect of water potential on changes in fresh mass of shoots during germination. Fresh mass of shoot (mg) Germination (days) Gupta et al .(1991) Ludhiana 2 4 6 2 4 6 Control stress Control stress : 0 MPa Control, : -0.14MPa stress

Table 11 : Amount of seed Carbon (mg) at maturity remobilised and assimilated during podding:

36 Table 11 : Amount of seed Carbon (mg) at maturity remobilised and assimilated during podding Genotype Seed yield (kg/ha) Treatment Post- podding Pre- podding Total (C) Assimilated (C) Remobilised (C) Tyson 7.3 Control 3058 302 3360 4.1 Stress 1508 292 1800 Kaniva 5.3 Control 2130 154 2284 1.5 stress 574 52 626 Australia Davies et al . (1998) 36

Table 12: Amount of seed N (mg) at maturity remobilised and N uptake during podding. :

37 Table 12: Amount of seed N (mg) at maturity remobilised and N uptake during podding. Genotype Seed yield (kg/ha) Treatment Post- podding Pre- podding Total (N) Uptake (N) Remobilised (N) Tyson 7.3 Control 30 170 200 4.1 Stress 4 151 155 Kaniva 5.3 Control 56 92 148 1.5 stress 5 48 53 Australia Davies et al . (1998) 37

Table 13: Varietal response of seed yield in irrigated and stress conditions. :

38 Table 13: Varietal response of seed yield in irrigated and stress conditions. Sr. No Genotypes Seed yield/plant (g) % decrease in yield in E2 Rank Irrigated (E1) Stress (E2) 1. K-850 8.0* 3.5* 55.5 3 2. GICG-9228 7.4 3.2* 57.0 4 3. KPG-316 7.3 3.1* 57.0 5 4. ICCV-10 7.3 4.2* 42.0 1 5. BG-1032 6.1 3.2* 48.0 2 6. Annegeri-1 ( C ) 4.8 1.8 62.1 - S.Em  0.5 0.1 C.D. (P=0.01) 2.9 0.9 C.V.% 11.2 7.5 Bangalore Jagannath et al. (1999) 38

Table 14: Effect of terminal drought on seed yield production in rainfed and irrigated condition.:

39 Table 14: Effect of terminal drought on seed yield production in rainfed and irrigated condition. Genotypes Seed yield (g/plant -1 ) Irrigated Rainfed Mean C-214 5.7 5.5 ( 2.9 ) 5.6 K-850 6.2 5.9(5.7) 6.1 Annegeri 6.2 5.9(4.3) 6.0 BG-362 6.4 5.1 (20.9) 5.7 H-208 5.2 4.6 (11.3) 4.9 Amethyst 5.2 4.7 (10.4) 4.9 Tyson 5.1 4.0( 21.0 ) 4.5 Genotypes Treatments - S.Em  0.214 0.121 - CD at 5% 0.614 0.347 - Value in parenthesis indicates per cent decrease. New Delhi Kumar et al. (2001) 39

Table 15: Effect of terminal drought on harvest index on chickpea genotypes:

40 Table 15: Effect of terminal drought on harvest index on chickpea genotypes Genotypes Harvest index (%) Irrigated Rainfed Mean C-214 39.88 44.57(11.76) 43.23 K-850 42.12 44.29( 5.1 ) 43.21 Annegeri 38.38 43.39(13.03) 40.88 BG-362 34.11 38.91(14.07) 36.51 H-208 32.44 40.44( 24.66 ) 36.44 Amethyst 33.29 39.67(19.16) 36.48 Tyson 35.17 38.41(9.2) 36.79 Genotypes Treatments G x T S.Em  1.236 0.618 1.748 CD at 5% 3.570 1.785 NS Value in parenthesis indicates per cent increase New Delhi Kumar et al. (2001) 40

Table 16: Performance of ICCV-96029 and long duration control at two locations in India:

41 Table 16: Performance of ICCV-96029 and long duration control at two locations in India Characters Patancheru 1 Hissar 2 ICCV-96029 C-235 ICCV-96029 Pant G-114 Days to first flower 24  1.0 61  0.5 43  2.0 83  3.0 Days to first pod 29  0.5 69  0.0 75  4.0 107  4.0 Days to maturity 79  1.0 109  3.5 128  3.0 155  2.0 Reprod. phase (days) 55  2.0 48  4.0 85  4.0 72  5.0 Plant height (cm) 40  1.5 46  6.0 54  2.0 45  6.0 Seed yield plant -1 14  5.6 21  4.7 17  2.0 26  4.0 Seed yield kg ha -1 1022  84.0 1439  222.5 1042  58.0 2049  166.0 ICRISAT & Hisar Kumar et al. (2001) 41 1 Mean of two environment 2 Mean of three environment

Table 17 : Mean, range PCV(%), GCV(%), Heritability and genetic advance in rainfed and irrigated conditions :

42 Table 17 : Mean, range PCV(%), GCV(%), Heritability and genetic advance in rainfed and irrigated conditions 42 Characters Mean Range PCV(%) GCV (%) Heritability Genetic advance Days to 50% flowering R 54.1 I 53.9 42.0- 63.5 44.0- 63.5 11.2 10.26 10.5 9.8 89.2 91.7 20.6 19.4 Days to maturity R 94.6 I 86.3 87- 102.5 81.5- 91.0 4.8 3.5 3.7 2.7 59.7 59.2 5.9 4.3 No. of primary branches R 4.7 I 4.7 2.8- 6.4 3.6- 6.10 13.4 11.9 13.1 13.0 50.4 54.6 19.2 19.8 No. of secondary branches R 9.3 I 5.3 6.7- 13.6 2.7-7.15 18.4 17.6 16.2 22.1 85.7 85.5 31.0 42.2 No.Pods /plant R 19.6 I 8.0 15.7-26.0 4.3-11.45 16.0 22.2 14.1 18.4 77.9 68.8 25.8 31.5 Shelling (%) R 74.0 I 71.2 61.0-82.4 61.7-79.5 8.4 5.9 6.5 5.0 60.0 72.6 10.3 8.9 100-seed weight (g) R 23.0 I 22.7 13.2-34.1 13.3-32.1 29.6 27.6 28.8 26.5 94.6 55.0 57.7 31.3 Seed yield /plant (kg/ha) R 5.7 I 2.8 3.4-8.0 1.1-3.78 24.8 32.4 22.1 31.5 79.5 94.6 40.7 63.2 Harvest index(%) R 49.7 I 35.0 37.7-67.6 24.4-46.3 14.1 15.7 12.7 13.9 81.5 77.5 23.7 25.7 Biomass on 46 th day R 8.5 I 6.4 4.6-13.5 4.4-12.4 29.2 28.9 25.1 25.6 74.0 78.1 44.5 46.6 Biomass on 76 th day R 16.6 I 9.6 32.0-6.2 18.5-30.1 30.1 25.93 28.2 24.1 91.8 86.3 57.7 46.1 Biomass accumulated during stress period R 8.0 I 3.4 2.7-17.2 1.65-9.8 45.8 45.2 43.9 44.2 91.8 95.4 86.9 89.0 Bangalore Jagannath et al .(1999) R = Rainfed, I = Irrigated

Table 18: The estimates of genetic parameters of variability for seed yield and other traits in chickpea under rainfed and irrigated conditions.:

43 Table 18: The estimates of genetic parameters of variability for seed yield and other traits in chickpea under rainfed and irrigated conditions . Sr. No. Character Range GCV PCV Heritability (BS%) Expected GA as % mean 1. Days to 50% flowering R 54.00- 75.6 I 55.00- 80.3 6.2 7.3 6.5 7.5 92.0 95.7 12.3 14.8 2. Days to maturity R 108.6- 122.00 I 122.3- 130.3 3.2 1.5 3.3 1.7 91.8 78.1 6.4 2.8 3. Plant height (cm) R 37.5- 57.5 I 53.3- 74.7 8.5 6.7 8.9 7.0 91.8 92,8 16.8 13.4 4. Plant spread (cm) R 12.7- 23.8 I 14.6- 29.0 11.4 11.2 12.7 12.1 81.1 86.3 231.2 21.5 5. No. of basal branches/plant R 1.80- 3.40 I 1.9- 3.3 11.8 10.0 13.6 12.6 74.1 66.9 21.0 16.5 6. No. of secondary branches/plant R 11.2- 19.6 I 11.0- 19.6 12.7 10.5 14.4 12.6 77.7 69.3 23.1 18.1 7. Pods/plant R 36.1- 132.6 I 51.80- 120.6 18.6 12.1 22.3 16.2 69.7 55.8 32.0 18.7 8. Seeds/pod R 1.0- 1.9 I 1.2- 2.0 16.9 11.4 20.0 14.8 70.0 62.9 29.4 18.3 9. 100- seed wt. (gm) R 10.8- 34.6 I 13.0- 38.7 22.0 22.9 22.2 23.2 98.1 98.1 44.9 46.8 10. Seed yield/plant (gm) R 7.0- 21.7 I 13.64- 27.1 19.0 17.0 22.5 19.1 71.6 79.3 33.2 31.2 11. Protein content (%) R 18.0- 23.9 I 18.0- 25.3 5.9 7.1 8.1 9.4 51.9 57.6 8.7 11.1 Shinde et al. (1996) Rahuri R = Rainfed, I = Irrigated

Table 19 : Genetic Parameters on different traits under rainfed conditions:

44 Table 19 : Genetic Parameters on different traits under rainfed conditions Sr. No. Characters Mean Range PCV GCV 1. Days to maturity 139 133 -144 2.1 2.03 2. Plant height (cm) 44.0 33.4 - 51.4 10.7 10.1 3. Pods/plant 113 71- 141 15.1 14.8 4. Biological yield (g) 71.1 53.0 -99.8 17.0 16.7 5. Seed yield /plant (g) 30.1 18.5 -46.8 25.5 25.2 6. 100- seed weight (g) 18.1 11.9 -32.4 30.6 30.6 7. Membrane injury (%) 59.8 45.4 -74.4 15.0 15.7 8. Relative water content (%) 69.9 62.8 - 81.6 7.6 7.3 Meena et al. (2006) New Delhi 44

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45 45 SOURCE OF RESISTANCE

Table 20: Annual wild species of chickpea and their importance. :

46 Table 20: Annual wild species of chickpea and their importance. 46 Species Somatic chromosome number Source of resistance Cicer arietinum 14,16,24, 28,33,32 Ascochyta blight, f usarium wilt, drought , cold, leaf minor Cicer cuneatum 16 Leaf minor, ascochyta blight Cicer judaicum 16 Leaf minor, cold Cicer pinnatifolium 16 Leaf minor, cold Cicer reticulatum 16 Fusarium, cold , blight Cicer bijugum 16 Blight, cold cyst nematode Cicer echinospermum 16 Leaf minor, cold Chopra (2000) New Delhi

Table 21: Sources of single and multiple resistance to abiotic and biotic stresses in chickpea identified at ICARDA and ICRISAT.:

47 Table 21: Sources of single and multiple resistance to abiotic and biotic stresses in chickpea identified at ICARDA and ICRISAT. 47 Stress Source of resistance Single stress Ascochyata blight ILC 72, 200, 3279, 3856, 5902,6090 ; ICC 1069 Botrytis grey mold ICC 1084, 1102, 3540, 4065 Fusarium wilt ILC 54, 240, 256, 336, 487, WR 315 ; ICC 11550, 12467 Pod borer ICC 506, 6663, 10619, 10667; ICCV 7 Cold ILC 8262, 482 M Drought ICC 4958 Multiple disease stress Wilt/ dry root root rot/black root rot ICC 12237, 12269 Wilt /dry root rot ICC 11315, 12241, 14372, 12275 Wilt/stunt virus ICC 10136, 10805, 11502, 11551 Singh and Saxena (1993) Syria

Table 22: Inheritance of different traits used as indicators of drought resistance:

48 Table 22: Inheritance of different traits used as indicators of drought resistance 48 Characteristics Crop Genetics Reference Dehydration avoidance Glabrous leaves Soybean Single dominant gene Singh (1983) Stay leaves green Cowpea Rds-1, Rds-2 gene, monogenic resistance Kodomi et al. (1999) Early flowering Chickpea efl 1 gene monogenic resistance Anbessa et al. (2006) Dehydration tolerance Pod formation and no flower abscission Rajma Partially dominant gene Singh (1983) Proline accumulation Brassica Additive gene action Singh (1983)

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BREEDING APPROACHES

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50 1. Conventional methods Introduction Selection Hybridization Pedigree method Backcross method d. Mutation breeding 2 . Nonconventional methods Genetic transformation Marker assisted selection 50

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51 Figure 7: Breeding approach to develop varieties suitable for a range of environment. Singh (1983) New Delhi

Figure 8: Breeding approach to for drought resistance based on combined use of optimum and moisture stress environment.:

Figure 8: Breeding approach to for drought resistance based on combined use of optimum and moisture stress environment. Singh (1983) New Delhi

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53 Achievements 53

Table 23: Drought tolerant varieties developed at ICRISAT, Hyderabad:

54 Table 23: Drought tolerant varieties developed at ICRISAT, Hyderabad Sr.No . Variety No. Days to flower Days to maturity Yield (kg/ha) 100 seed weight (g) 1. ICCV 98944 44 101 2153 14.05 2. ICCV 98939 50 91 2136 11.45 3. ICCV 98936 48 96 2131 11.15 4. ICCV 98903 40 88 2038 28.15 5. ICCV 98902 41 89 1995 29.65 6. ICCV 98932 49 96 1977 10.85 7. ICCV 98905 42 88 1952 20.1 8. ICCV 98921 49 93 1916 11.1 9. ICCV 98904 42 80 1900 25.8 10. ICCV 94924-2 36 83 1898 14.65 11. ICCV 98928 43 91 1888 10.15 12. ICCV 98908 47 95 1872 15.4 13. ICCV 94916-4 41 96 1864 30.1 14. ICCV 98947 42 87 1862 10.25 ICRISAT Anonymous (2004) 54

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Biotechnological approaches

Gene Expression :

56 Gene Expression The molecular studies based on dehydration stress are mainly desication tolerance to maturing embryo and resurrection based on Arabidobsis thaliana and other crops such as tomato, pea, wheat and barley. Genes regulated by drought stress can be divided in to three groups. Gene encoding polypeptides of unknown function Gene encoding LEA (late embryogenesis abundant) protein and related polypeptides Gene encoding polypeptides of known function DNA sequence analysis of osmotic stress inducible cDNAs indicate that genes responsible for drought encode variety of proteins. Many of these proteins encoded by these cDNAs have been classified in to various groups LEA (late embryogenesis abundant) RAB (responsive to ABA) or dehydrin proteins The LEA proteins, first identified during seed maturation and desiccation, express in water stressed vegetative tissues in almost all plants. They protect the dehydrating cells by a variety of mechanisms, including renaturation of unfolded protein. 56

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57 57 Water channel proteins in plants which related to super family of membrane intrinsic proteins(MIPs) first characterized in Escherichia coli. Several MIP- related proteins have been identified in plants. Trg 31 in pea is initially identified when gene expression in induced in partially dehydrated leaves. NOD 26 in Arabidobsis thaliana dehydration inducible genes identified . Vsp in soyabean is vegetative storage protein

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58 Glycinebetaine is one of the compatible solutes that are well established to protect key cellular metabolic events against damage. The codA gene for choline oxidase (the enzyme that converts choline into glycinebetaine) isolated from Arthrobacter globiformis could be successfully introduced into chickpea through Agrobacterium mediated transformation following the perfect protocol that was fine-tuned earlier . Presence and expression of the codA gene in the T 3 population was confirmed through PCR, Western analysis and activity of choline oxidase. Most importantly, the codA transgenics showed the presence of glycinebetaine, while no traces of glycinebetaine were found in wild type. No significant alteration in the levels of choline, which is the substrate used by choline oxidase for synthesis of glycinebetaine, was identified in the codA transgenics in comparison with wild type. Development of transgenic chickpea with enhanced tolerance to drought stress using the CodA gene 58 Sharma (2007) ICRISAT , Hyderabad

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59 Figure 9: A group of codA transgenic genotypes of chickpea grown during off-season. 59

Transgenic chickpea for tolerance to drought:

60 Transgenic chickpea for tolerance to drought Arabidopsis thaliana plants P5CSF129A gene or rd29A gene Agrobacterium mediated , CaMV 35S promoter encoding pyrrolin-5-carboxylate synthetase inducing tolerance to drought stress in chickpea 60 Sharma (2007) ICRISAT , Hyderabad

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61 Figure 10: Screening of transgenic chickpea events transformed with 35S:P5CSF129A or rd29A:DREB1A genes for inducing tolerance to drought stress . 61

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62 62 Conclusions: Chickpea ( Cicer arietinum ) is the third most important grain legume in the world, grown extensively in all continents especially in semi- arid tropics under rainfed ecosystem. Chickpea being rainfed crop, significant improvement in yield was achieved through minimizing the impact of biotic and abiotic stresses through incorporation of tolerance or resistance Among the various abiotic stresses affecting chickpea, drought is the most important constraint causing considerable production losses . Lack of simple repeatable screening technique has been the major hindrance for breeding chickpea for drought tolerance. Large number of morphological and physiological traits contribute to drought tolerance or resistance; improvement in chickpea is possible only if all the traits are considered in conjunction though conventional and biotechnological approach such as gene pyramiding or stacking.

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63 Consolidation of yield gains through multiple resistance to various abiotic and biotic stresses. Efforts to be made for designing varieties with determinate growth habit and efficient source-sink kinetics addressing the physiological constraints. To develop a plant ideotype by restructuring the morphological attributes that can withstand drought conditions. Broadening the genetic base through wide hybridization. Development of a more saturated and well distributed genome map so as to identify the molecular markers and tagging the genes of interest Incorporation of molecular breeding or marker assisted selection of breeding material. 63 Future strategies :

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64 64 Thank You

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