heterosis, combining ability and gene action in pigeonpea

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HETEROSIS, COMBINING ABILITY AND GENE ACTION IN PIGEONPEA

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1 Presented By:- Arun D. Ugale M. Sc. (Agri .) Genetics And Plant Breeding HETEROSIS, COMBINING ABILITY AND GENE ACTION IN PIGEONPEA

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FLOW OF PRESENTATION INTRODUCTION BASIC REQUIRMENT FOR HYBRID VARIETIES HETEROSIS COMBINIG ABILITY GENE ACTION CONCLUSION 2

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India is self-sufficient in food grain production . There is need to enhance the total pulse production to provide sufficient amount of food to the increasing population. Majority of people in India are vegetarian. Pulses add atmospheric nitrogen in the soil besides meeting their own requirement of nitrogen. Pigeonpea ( Cajanus cajan (L.) Millsp ) is second most important pulse crop grown in India after chickpea which contain 20-25 per cent protein which is nutritionally, complementary to cereals in its pattern and profile of amino acids. 3

Area, production and productivity: 

Area, production and productivity INDIA GUJARAT AREA 35,00000 ha PRODUCTION 24,00,000 MT PRODUCTIVITY 685 Kg/ ha AREA 26600 ha PRODUCTION 2627 MT PRODUCTIVITY 987 kg/ha 4 Gandhinagar Anonymous, 2008 Singh, 2008 Kanpur

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Table 1. Nutritional c onstituents of pigeonpea Constitute Green seed Mature seed Protein (%) 21.0 18.8 Starch (%) 44.8 53.0 Soluble sugars (%) 5.1 3.1 Crude fiber (%) 8.2 6.6 Fat (%) 2.3 1.9 Calcium(mg/100g) 94.6 120.8 Magnesium (mg/100g) 113.7 122.0 Copper (mg/100g) 1.4 1.3 Iron (mg/100g) 4.6 3.9 Zinc (mg/100g) 2.5 2.3 5

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FOUR BASIC REQUIREMENT FOR DEVELOPMENT OF HYBRID VARIETIES Efficient systems for genetic emasculation Sufficient amount of out crossing for pollen transfer Easy and cheap production of quality hybrid seeds Exploitation of heterosis/hybrid vigour 6

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1. EFFICIENT SYSTEMS FOR GENETIC EMASCULATION eg. GMS, CMS, CGMS, self incompatibility and pistillate lines 7

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GENETIC MALE STERILITY In pigeon pea, two types of genetic male sterility, each contribute by single recessive gene has been reported. A) Reddy et al. (1978) identified a white translucent male sterile anther. Six male sterile plants were identified at ICRISAT in 1974, derived from totally two unrelated species. 8 ms-3A } From ICP-1555 ms-3B ms-3C ms-4D ms-5E ms-4A from ICP-1596

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Both male sterile lines are medium in maturity (180-200 days). Non determinate branching pattern. Seed colour of ms-3A group is cream while ms-4A group is brown. Male sterile plants are morphologically similar to normal plants except senescence is smaller than fertile plants. All the floral parts except anthers are smaller in size and colour in male sterile as well as male fertile plants. Anthers of ms plants are smaller, flat and translucent with whitish scaly surface, no dehisce, dry faster than normal anthers. CHARACTERISTICS OF MALE STERILE LINE 9

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Male sterility is governs by a single recessive gene. Maintenance of male sterility is done by sibbing male sterile and fertile plants observed in the same population. The visible marker, i.e. White translucent anther provides an efficient way of recognizing these ms plants in the field. 10 ms ms Ms ms . . ms ms Ms ms Expelled out Male Sterile Male Fertile Male Sterile Male Fertile

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11 B) Saxena et al . (1983) reported arrow head shaped brown colour male sterile anthers. i.e. A new source of Genetic male sterility has been reported from Australia (Wallis et al. 1981) . Anthers of this male sterile source are brown , shriveled, non-dehiscent and arrow shaped and contained no pollen grains. e.g. ms prabhat DT; ms prabhat NDT; Gms-1,2,7; T-15-15 etc. Saxena et al. (1989) reported that both male sterile and female fertile are being used in hybrid breeding programme in pigeon pea.

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GMS BASED WORLD’S FIRST HYBRID ICPH-8 MS. P. DT x ICPL 161 12

ICPH-8: 

ICPH-8 Characteristics Female Male Hybrid Name MS. P. DT ICPL 161 ICPH 8 Type Determinate Indeterminate Indeterminate Growth habit Errect Semi spreading Semi spreading Plant height (cm) 90-120 150-180 160-180 Days to flowering 60-70 70-75 75-80 Days to maturity 110-120 130-135 130-140 Flower colour Yellow Yellow Yellow Grain colour Red Red Red Grain size Medium Medium Medium 13 The first hybrid pigeon pea ICPH-8 was developed by ICRISAT by using genetic male sterility and released for cultivation in India in 1990.

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Two approaches were adopted to achieve this goal. Wide hybridization Mutagenesis 14 HOW TO INDUCE CYTOPLASMIC MALE STERILITY ?

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A. Hybrid involving wild Cajanus sp as female parent Cajanus scrabacoides Cajanus sericeus Cajanus cajanifolius Cajanus albicans B. Hybrid involving pigeon pea as female parent Cajanus acuntifolius 15 CGMS was reported to occur in many wild species of pigeonpea. This has been successfully achieved by using wild relatives through repeated backcrossing . Wide hybridization

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2) Mutagenesis Cytoplasmic mutations caused by certain chemicals and physical mutagens leading to CMS in plants are reported in several crop species. These mutagens have been claimed to act specifically on DNA containing cytoplasmic inclusion like mitochondria and chloroplast inducing CMS mutation (Zaveri, 1992.) 16

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CYTOPLASMIC MALE STERILITY The first breakthrough in this technology was achieved when a CMS system was developed by crossing C. cajanifolius (a wild species) and a cultivated line. This CMS system is stable across diverse environments and has an excellent fertility restoration system. After the development of a stable CMS system, several experimental hybrids were produced and evaluated. Kandalkar (2007) found that CMS-based hybrids recorded standard heterosis up to 156% for grain yield. Saxena (2007) reported yield advantage of 50 to 100% over the popular varieties/checks. 17 17

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ICPH 2671 The new hybrids developed on an improved A-line (ICPA 2043) showed promise with respect to yield, seed size and disease resistance. Important characteristics of hybrid ICPH 2671 Growth habit:- Semi-spreading and indeterminate Days to flowering:- 114–120 days Days to 75% pod maturity:- 166-184 days Petals:- Yellow with dark red streaks Pods:- Dark purple in colour Photo-sensitive Highly resistant to Fusarium wilt and sterility mosaic diseases. 18

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CGMS BASED GTH- 1 : (CMS GT288A x GTR 11) Worlds first CGMS based hybrid of pigeonpea developed by the SDAU by utilizing cytoplasm of wild species of Cajanus ( Cajanus scarabacoides ). This has resulted in the identification of stable CGMS lines (A/B). Subsequently fertility restoration mechanism was also perfected from two sources viz., Fertile advanced lines of Cajanus scarabacoides x Cajanus cajan Pigeonpea germplasm sources 19

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Early maturity [ 150 + 5 days) Grain yield 32 and 42% higher than GT 101 and AKPH - 4101, respectively Non determinate growth habit Green pod with streaks at base Bold white seed Better dal recovery [87.2%] Yield potential: 2827 kg/ha 20

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Pigeonpea is especially self pollinated crop due to their flower structure but out crossing may result from bee activities (Pathak ,1970) The percent out crossing has been reported from 3% to 40% (Khan,1973). Very high degree of out crossing up to 94.5% has been reported from Kenya. Saxena et al. (1989) reported 5 to 70% out crossing in pigeonpea. 21 2) SUFFICIENT AMOUNT OF OUTCROSSING FOR POLLEN TRANSFER

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Flowering habit of varieties Nature of glumes Colour of petals Number of insect pollinators Location of the field in relation with insect habitat Climatic condition Factor affecting natural out crossing in pigeon pea [ Bhatia et al. (1981) ] 22

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3) Easy and cheap production of quality hybrid seeds 23 Seed production of male sterile lines:- The genetic male sterility in pigeonpea is governed by a pair of single recessive gene (ms ms). Therefore, it is maintained in heterozygote. For seed production of male sterile line the sibbed seeds (Ms ms x ms ms) are planted in isolation. This population is segregate into 1:1 sterile and fertile plants. All the plants are identified and only sterile plants are tagged and fertile plants are retained without tagging. The male sterile plants are pollinated naturally by pollen from the fertile plants (Ms ms) by the insect pollinator.

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24 B) Seed production of pollinator line: The seed of pollen parent can be produced easily in isolation. Besides, the seed of the pollen parent from the hybrid seed production block may be harvested without any additional investment.

PLAN OF HYBRID SEED PRODUCTION: 

PLAN OF HYBRID SEED PRODUCTION Parents Female :- CMS GT288A Male :- GTR 11 Seed rate Female :- 25 kg/ha Male :- 5 kg/ha Spacing Female :- 60 X 10 Male :- 60 X 20 25

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* I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * * I I I I I * I I I I I * 5:1 1 7 13 LAYOUT OF HYBRID SEED PRODUCTION **** Restorer Line ---- Female Line 26 * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * * I I I I * I I I I * 4:1 1 6 11

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EXPLOITATION OF HETROSIS 27

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The term heterosis was first used by Shull in 1914 . Heterosis refers to the superiority of F1 hybrid in one or more characters over its parents. The term hybrid vigour is used as synonym for heterosis. Heterosis differs from luxuriance. The former refers to increase of F1 over parents. in general vigour ,yield and adaptation ,where as later refer to increase of F1 over parents in vegetative growth but not yield and adaptation . INTRODUCTION 28

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1 ) Superiority over parents 2 ) Confined to F1 3 ) Genetic control 4 ) Reproducible 5 ) Associate with SCA 6 ) Effect of heterozygosity 7 ) Conceals recessive genes 8 ) Low frequency 29 IMPORTANT FEATURE OF HETEROSIS

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ESTIMATION OF HETROSIS 1 ) Average heterosis. When the heterosis is estimated over the mid parent, i.e, mean value or average of the two parents, it is known as average heterosis, which is estimated as follows : Average heterosis = Where, F 1 = the mean value of F 1 MP= the mean value of two parents involved in the cross 30

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2) Hetrobeltiosis When the heterosis is estimated over the superior or better parent, it is referred to as hetrobeltiosis. It is worked out as follows Hetrobeltiosis = Where, BP= mean value (over replication) of the better parents of the particular cross 31

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32 3) Useful heterosis The term useful heterosis coined by Meredith and Bridge (1972). It refers to the superiority of F 1 over the standard commercial check variety. It is also called as economic heterosis. It is estimated as follows. Useful heterosis = Where, CC = mean value over replications of the local commercial cultivar

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Cross Plant height Primary branches per plant 100 seed weight Harvest index UPAS 120 X ICPL84023 31.54** 2.40 -2.69 -1.41 ICPL 84023 X KVBR 80-2-1 82.26** -11.93 -6.21 -2.18 ICPL 87119 X KVBR 80-2-1 1.59 -11.99 -6.34 -7.08 ICPL 90035 X KVBR 80-2-1 -3.97 -9.92 -2.06 29.44** *,** Significant at 5% and 1% level of significance, respectively BP= Better parent; MP= Mid parent Table 2. Heterobeltiosis in four pigeonpea crosses Pantnagar Gupta et al. (1996) 33

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Cross combinations Days to initial flowering Plant height (cm) Primary branches per plant Secondary Branches per plant KPMS 1050 x DA 94-6 4.02* -9.96** 10.59 91.30** KPMS1050 x Bahar meant 6.32** -16.47** 22.73** 160.87** KPMS 1050 x Pusa-9 4.31* -18.86** 31.82** 137.70** DAMS-1 x Sharad 1.72 -13.02** 31.82** 205.78** KPMS1050 x DA 34 3.73 -17.40** 22.73** 165.22** DAMS-1 x DA 93-1 2.30 -13.42** 24.23** 88.39* DAMS-1 x DA 94-6 2.30 -17.26** 40.90** 169.56** KPMS1050 x DA 924 : 5.46** -5.45 28.77** 76.82* KPMS1050 x DA 37 2.59 -13.15** 40.90** 110.13** DAMS-1 x DA 93-5 0.86 -15.14** 21.23** 108.70** * Significant at P = 0.05 ** Significant at P = 0.01. Table 3:- Standard heterosis for eight characters in long duration pigeon pea hybrids ever Bahar Pusa Pandey (2004) 34

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Cross combinations Number of clusters/ plant Number of pods/ plant Harvest index Seed yield per plant KPMS 1050 x DA 94-6 89.06** 50.18** 90.01** 144.32** KPMS1050 x Bahar meant 88.68** 55.23** 41.16 121.12** KPMS 1050 x Pusa-9 94.03** 45.13* 28.14 113. 90** DAMS-1 x Sharad 100.25** 69.50** 28.66 112.36** KPMS1050 x DA 34 113.05** 103.64** -8.42 109.79** DAMS-1 x DA 93-1 75.37** 41.84* 37 19 99.47** DAMS-1 x DA 94-6 96.27** 46.63** 27.82 96.38** KPMS1050 x DA 924 : 55.10* 44.42* 15.06 90.70** KPMS1050 x DA 37 55.34* 49.56** 51.78* 89.16** DAMS-1 x DA 93-5 85.20** 48.85** 49.06* 86.59** * Significant at P = 0.05 ** Significant at P = 0.01. Table 3 cont……… Pusa Pandey (2004) 35

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Charcter Range of hetrosis Best hetrotic cross over standard check hetrobeltiosis Standard hetrosis Days to flowering -5.71-22.22 -9.51-20.55 MS Prabhat (DT) x 12-33 Days to maturity 12.17-12.27 -18.48-2.46 QMS-1 x Sel 90311 MS Prabhat (DT) x 12-33 Plant height -29.12-32.59 -32.89-33.00 QMS-1 x P 610 Plant spread -30.80-50.48 -25.06-84.10 QMS-1 x sel 90307 Clusters/plant -35.45-111.21 -71.63-97.40 MS Prabhat (DT) x sel 90309 Pods/clusters -36.50-93.24 -41.59-129.20 MS Prabhat (NDT) x sel 90311 Pods/plant -26.56-131.78 -33.37-6.98 QMS-1 x sel 90311 MS Prabhat (NDT) x P 610 ( Seeds/pod -1.38-18.66 -12.24-11.46 QMS-1 x sel 90309 100- seed weight -31.69-25.49 -29.3-22.93 MS Prabhat (NDT) x sel 90312 Seed yield/ plant -23.57- 171.58 -36.19- 51.31 QMS-1 x sel 90307 (51.31 % ) QMS-1 x sel 90311 (50.90%) QMS-1 x sel 90310 (48.44%) QMS-1 x sel 90309 (47.30%) MS Prabhat (NDT) x sel 90306 (40.0%) Biomass/plant -41.63-211.11 -53.62-56.57 MS Prabhat (NDT) x H 87-2 Harvest index -36.87-26.49 -35.62-31.23 QMS-1 x P 610 Figures in paranthesis denote standard hetrosis values, *,** Significant at 5% and 1% level,respectively Table 4 . Range of heterosis and most hetrotic crosses for twelve characters in pigeonpea New delhi Reddy et al . (2004) 36

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Table 5:- Analysis of variance for seed yield per plant and its components in pigeon pea Source df Seed yield per plant (g) Days to 50% flowering Days to maturity Plant height (cm) No. of branches per plant No. of pods per plant Replication 2 6.57 85.50 246.21 364.45 2.87 383.22 Parents (P) 8 1125.49** 1175.49** 2043.44** 3608.92** 31.00** 6265.55** Hybrids (H) 35 3196.16** 481.20** 728.86** 811.52** 36.11** 65432.14** P V/s H 1 19203.01** 1229.59** 3686.66** 6324.64** 248.10** 511940.32** Error 160 27.95 91.92 95.51 180.82 1.52 606.88 Sardarkrushinagar Yadav et al. (2005) 37 Source Pod length (cm) Seeds per pod 100- seed weight Harvest index (%) Shelling percentage Percent pod damage Replication 0.11 0.01 0.71 4.17 31.67** 4.44 Parents (P) 3.60** 2.90** 7.25** 762.02** 547.49** 166.45** Hybrids (H) 3.37** 0.67** 9.69** 719.94** 545.67** 355.46 P V/s H 1.69* 0.19 0.76 878.67** 483.87** 211.43** Error 0.34 0.13 1.06 4.83 4.81 4.38 *,** Significant at 5 and 1 percent level respectively

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Character Range of heterosis over Number of crosses showing significant desirable heterosis BP SP BP SP Seed yield per plant (g) -89.22 - 601.28 -93.33 - 147.62 30 19 Days to 50 per cent flowering -10.94 - 14.87 30.21 - 40.23 3 0 Days to maturity -11.34 - 12.14 -1.03 - 31.48 7 0 Plant height (cm) 0.08 - 93.75 46.42 - 140.67 0 0 No. of branches per plant -34.59 - 139.24 -56.07 - 100.54 27 5 No. of pods per plant -74.27 - 697.80 -74.26 - 355.96 35 18 Pod length (cm) -35.23 - 45.70 -56.07 - 6.68 10 0 Seeds per pod -33.33 - 30.16 -16.98 - 46.03 13 0 100- seed weight (g) -33.80 - 27.76 -38.89 - 27.07 6 2 Harvest index (%) -86.84 - 316.79 -95.37 - 70.91 16 3 Shelling percentage -73.37 - 175.93 -79.92 - 2.40 18 0 Per cent pod damage -77.053 - 170.19 -58.82 - 256.12 21 4 Table 6. Range of heterosis over better parent (BP) and standard parent (SP) along with number of crosses showing significant heterosis in desired direction for various characters in pigeonpea. Sardarkrushinagar Yadav et al. (2005) 38

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Promising hybrids Seed yield per plant (g) Standard heterosis Significant desirable standard heterosis for component traits GT-100 x ICP-12116 130.00 147.62** PP , HI Banas x ICP-9140 122.55 133.43** PP ICP-12161 x ICP-9135 114.50 117.33** PP GT-100 x ICP-12161 112.50 114.29** BP, PP GT-101 x ICP-12161 109.00 107.62** BP, PP , PPD GT-101 x ICP-12116 101.25 92.86** PP , HI GT-100 x Banas 94.63 80.33** PP GT-100 x ICP-11912 90.00 71.42** PP ICP-13555 X ICP-9135 89.27 70.04** PP , PL, PPD Banas X ICP-11488 80.33 52.95** PP , PPD * and ** significant at 5 % and 1 % levels, respectively. Where, BP= number of branches per plant. PP= number of pods per plant, PL= pod length, Hl= harvest index and PPD= per cent pod damage . Table 7. Promising hybrids for seed yield per plant with standard heterosis for major component traits showing significant desirable standard heterosis in pigeon pea. Sardarkrushinagar Yadav et al. (2005) 39

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Characters Range of heterosis over Best cross No. of significant crosses in desirable direction BP SC BP SC BP SC Days to 50% flowering -5.35 -42.61 -17.63-21.79 ICPL 85063 x ICP 8863 ICPL 85034 x ICP 8863 1 7 Days to maturity -5.67-23.10 -11.02- 12.20 ICPL 85063 x ICP 8863 ICPL 85034 x ICP 8863 1 15 Plant height 0.83- 25.29 -0.06-21.36 ICPL 85034 x ICPL 89044 LRG 30 x ICPL 87119 6 5 No. of primary branches per plant -9.33-24.40 -9.57-40.43 PRG-100 x ICP 8863 LRG 30 x ICPL 87119 5 9 No. of clusters per plant -37.69-19.35 -33.53-15.67 LRG 38 x ICP 89044 PRG-100 x ICPL 87119 3 4 No. of pods per plant -16.39-50.13 -34.09-37.43 PRG-100 x ICP 8863 LRG-30 x ICP 8863 5 3 100- seed weight -21.48-5.17 -25.08-10.11 PRG-88 x ICPL 87119 PRG-100 x ICP 8863 0 2 Seed yield per plant -18.06-53.14 -25.44-51.38 LRG-30 x ICP 8863 LRG-30 x ICP 8863 5 5 ** Significant at 1 percent level, * Significant at 5 percent level Table 8 . Heterosis over Better parent (BP) and standard check (SC) in pigeon pea Tandur Kumar et al . (2009) 40

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COMBINING ABILITY 41

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The concept of combining ability as a measure of gene action was proposed by Sprague and Tatum in 1942 working on maize. Combining ability refers to the capacity or ability of a genotype to transmit superior performance to its crosses . The value of an inbred line depends on its ability to produce superior hybrids in combination with other inbreds. 42

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Combining ability analysis helps in the evaluation of inbreds in terms of their genetic value, and in the selection of suitable parents for hybridization. It also helps in the identification of superior cross combinations. which may be utilized for commercial exploitation of heterosis. For combing ability analysis, cross have to be made either in diallel, partial diallel or line x tester fashion. Combining ability estimates GCA and SCA effects which are based on first order statistics (mean values). It also provides information about the gene action involved in the expression of various quantitative characters and thus helps in deciding the breeding procedure for genetic improvement of such traits. 43 MAIN FEATURES

TYPES OF COMBINING ABILITY: 

TYPES OF COMBINING ABILITY

GENERAL COMBINING ABILITY: 

GENERAL COMBINING ABILITY The average performance of a strain or genotype in a series of hybrid combinations is termed as general combining ability . The GCA is estimated from half sib families. The GCA variance has positive correlation with narrow sense heritability. GCA attributed additive genetic variance and additive x additive interaction. The GCA helps in the selection of suitable parents for hybridization. 45

SPECIFIC COMBINING ABILITY: 

SPECIFIC COMBINING ABILITY The deviation of a particular cross from the general combining ability is known as specific combining ability. The SCA is estimated from full sib families. The SCA variance has positive association with heterosis or hybrid vigour. SCA attributed dominance genetic variance, dominance x dominance and additive x dominance types of interaction . The SCA helps in the identification of superior cross combinations for commercial exploitation of heterosis. 46

Estimation of combing ability: 

Estimation of combing ability Selection of genotypes Making single crosses Evaluation of material Biometrical analysis 47

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Source Cropping systems Df. Days to maturity Plant height Pods per plant Grain yield Harvet index Gca Sole crop 6 329.7** 2166.3** 12606.7** 875.9** 53.3** Intercrop in sorghum 635.5** 771.3** 903.9** 135.2** 22.3** Intercrop in pearl millet 401.1** 876.9** 3103.2** 527.4** 23.4** Sca Sole crop 21 32.1** 309.1** 2277.1** 217.6** 15.0** Intercrop in sorghum 41.6** 207.0** 178.7** 50.4** 40.3** Intercrop in pearl millet 94.0** 171.7** 210.5** 70.0** 27.5** Error Sole crop 27 1.5 4.1 25.1 2.5 0.7 Intercrop in sorghum 0.7 4.1 3.8 0.5 1.9 Intercrop in pearl millet 1.5 4.2 11.9 0.8 0.4 б 2 gca Sole crop 33.1 206.4 1147.7 73.1 4.4 Intercrop in sorghum 66.0 62.7 80.6 9.4 -2.0 Intercrop in pearl millet 34.1 78.3 321.4 51.1 -0.4 б 2 sca Sole crop 30.6 305.0 2252.0 215.1 14.4 Intercrop in sorghum 40.8 202.90 174.9 50.0 38.4 Intercrop in pearl millet 92.5 167.5 198.6 66.2 27.0 б 2 gca/ б 2 sca Sole crop 1.1 0.7 0.5 0.3 0.3 Intercrop in sorghum 1.6 0.3 0.5 5.3 -0.05 Intercrop in pearl millet 0.4 0.3 1.6 0.8 -0.02 ** Significant at P=0.01 Table 9. Combing ability analysis (mean squares) for yield and yield components under three cropping systems in pigeonpea Parbhani Khapre et al. (1993) 48

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Character Crosses Per se F 1 performance Sca effects Gca status of parents Performance of parents P1 P2 Days to maturity PL 8796 x Prabhat 126.5 -6.7** L x H 163.2 125.8 ICP 6997 x Prabhat 127.7 -6.0** L x H 158.5 125.8 Daithna Local x Prabhat 128.7 -0.5** H x H 145.5 125.8 4785-2- V2 x Prabhat 130.2 1.0 H x H 144.2 125.8 PBNA 54 x Prabhat 131.7 -3.9** L x H 161.0 125.8 Pods per plant PBNA 54 x BDN 2 216.5 28.2** H x H 155.0 160.7 Daithna Local x BDN 2 201.0 9.7** H x L 162.9 160.7 Daithna Local x Prabhat 189.0 36.8** H x H 162.9 111.9 BDN 2 x Prabhat 182.9 18.3** L x H 111.9 160.7 PL 8796 x BDN 2 172.6 20.1** L x H 103.9 160.7 Grain yield ICP 6997 x PBNA 54 74.9 17.6** H x H 45.7 47.9 ICP 6997 x BDN 2 68.6 6.3** H x H 45.7 54.8 PL 8796 x BDN 2 66.1 11.4** L x H 37.1 54.8 Daithna Local x Prabhat 69.9 0.4 H x H 47.1 54.8 PBNA 54 x BDN2 61.8 -0.5 H x H 47.9 54.8 ** Significant at P= 0.01 Table 10. Combing ability of the best five crosses of pigeonpea based on pooled per se performance, SCA and GCA effects of parents in sorghum Parbhani Khapre et al. (1993) 49

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Sources d.f Mean Squares Days to flowering Plant height Primary branches/ plant Pods/ plant Seeds/ pod 100- seed weight Yield/plant Replication 1 24.00 486.00** 0.24 1084.90 0.02 -0.08 146.70** Line 2 1512.40** 4794.89** 3.97 30245.30** 0.02 0.11 1377.20** Tester 8 45.87 430.00** 3.06 4562.80 0.10 0.23 152.8 Line X Tester 16 50.01 159.14 4.17* 4422.70** 0.08 0.64 301.4** Error 26 28.62 87.88 1.44 862.70 0.04 0.22 25.0 б 2 gca 60.76 204.42 0.06 1081.77 0.002 0.04 38.63 б 2 sca 10.70 35.62 1.14 1780.02 0.001 0.05 38.27 Average degree of dominance 0.42 0.41 4.36 1.28 2.34 1.12 1.89 *, ** Significant at P=0.05 and P=0.001 respectively Table 11. Analysis of variance for combining ability, variance and Average degree of dominance in short duration pigeon pea Kanpur Kumar et al. (2001) 50

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Sources Mean Squares Days to 50% flowering Pods/ plant 100- seed weight Yield/plant Lines ms Prabhat NDT 8.11** 45.99** -0.06 5.49** ms Prabhat DT 1.83** -13.28* -0.02 4.60** IMS-1 -9.94** -32.70** 0.09 -10.09** SE(+) Lines 0.86 4.70 0.08 0.80 Testers AL 31 0.50 -35.31** -0.27 2.18 ICPL 85010 1.00 -9.79 0.09 1.51 P 604 -1.00 24.34 0.19 2.91 P 605 0.83 -28.78** -0.06 -7.25** UPAS 120 1.67 45.34** -0.24 2.55 AF 98 0.83 -17.13 -0.09 1.23 ICPL 88001 0.17 16.21 -0.02 5.20** P 610 -6.83** 21.21* 0.08 1.58 H 87-24 2.83 -16.09 0.33** -9.90** SE(+) Testers 1.17 9.41 0.15 1.60 *, ** Significant at P=0.05 and P=0.001 respectively Table 12. Estimates of general combining ability effects of parents of short duration pigeon pea Kanpur Kumar et al. (2001) 51 51

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Crosses SCA effects Mean seed yield/plant (g) SCA effect of parents Significant response in other traits for SCA effects Female Male MS 288 x SKNP 9231 5.45 22.64 1.16 3.50 PP, RP, HI and PC MS 288 x SKNP 9219 4.09 19.87 1.16 2.10 DF, DM, PH, BP MS(P) DTx SKNP 9512 3.88 14.76 0.35 -2.00 DF, DM, PP ,SP end HI. MSPusa 33 x SKNP 9256 3.39 17.54 0.16 1.46 PP and PC T 21 x GAUT 90-2 2.77 13.44 -1.68 -0.18 BP, PP, PL, SP, SW, RP MS 288 x SKNP 9204-1 2.67 14.84 16 1.51 BP, PP ,SW and HI MST21xAL15 2.26 13.81 -1.68 -0.72 BP, PP , SW and PC. MS 288 x SKNP 96-3 2.23 15.46 1.16 -0.46 BP, PP, SP MS(P) DTx SKNP 9510 1.82 1426 0.35 -0.45 DF, PP ,,RP MS(P)DT x SPL 87 1.70 1192 0.36 2.75 HI, PP , RP, HI Table 13. Ten best specific combiners for seed yield/plant and their performance for other traits in pigeon pea. Sardarkrushinagar Pawar and Tikka(2003) DF= Days to flowering SP= Seeds per pod BP= Branches per plant DM= Days to maturity PH= Plant height PC= Protein content (%) PH= Plant height TW= Test weight (g) PP= Pods per plant RP= Reproductive phase HI= Harvest index SW= 100-Seed weight 52

Slide 53: 

Table 14. Analysis of variance for combining ability for ten characters in long duration pigeon pea Source of variation d.f. Days to initial flowering Day to maturity Plant height (cm) Primary branches per plant Secondary branches per plant Hybrids 35 16.58 107.52* 191.89 12.82 425.89* Lines 11 18.76 110.51* 270.76* 3.58 230.52** Testers 2 1.11 86.22 23.83 46.03** 3778.07** Line x Testers 22 16.90 107.96** 167.73 14.43 218.83 Errors 100 12.06 77.31 189.27 22.80 266.90 б 2 GCA -0.02 -0.03 1.52 -0.10 13.04 б 2 SCA 1.62 10.22 -7.18 -2.79 -16.02 б 2 GCA / б 2 SCA -0.01 -0.003 -0.21 0.04 0.82 *Significant at P = 0.05, **Significant at P = 0.01 Pusa Pandey (2004) 53

Slide 54: 

Source of variation d.f . Number of clusters per plant Number of pods per plant Per cent pod setting Harvest index Seed yield per plant Hybrids 35 28398.76** 27409.86** 3.79** 71.99* 2052.92* Lines 11 19127.23** 25333.08** 4.53* 55.64 1491.24* Testers 2 288768.15** 173053.56** 20.36** 3.20 14138.59** Line x Testers 22 9364.59 15207.92** 1.91 86.42* 1235.07 Errors 100 11418.46 13198.13 2.78 64.24 1245.46 б 2 GCA 1193.74 768.45 0.12 -0.91 51.50 б 2 SCA -684.63 669.93 -0.29 7.39 -3.46 б 2 GCA / б 2 SCA -1.75 1.15 -0.42 -0.13 -14.88 *Significant at P = 0.05, **Significant at P = 0.01. Table 14 Cont….. Pusa Pandey (2004) 54

Slide 55: 

Cross Combination Days to initial flowering Primary branches per plant Secondary branches per plant Number of clusters per plant Number of pods per plant Per cent pod setting Seed yield per plant KPMS1050 x Bahar mutant -0.19 -0.70 3.15 26.37** 51.74** -0.61 42.56** MS 3783 x Pusa-9 2.58 -1.08 24.99** 23.99** 53.00** 12.32** 41.47** KPMS 1050 x DA 34 -0.30 0.53 1.07 24.49* 59.26** -0.31 38.12** DAMS-1 x Bahar mutant 1.39 20.00** 28.82** 43.63** 44.29** -0.200 36.56** MS 3783 x DA 93-2 5.14* 12.92** 0.65 29.57** 28.55** 8.28** 36.47** MS 3783 x Bahar mutant -1.19 10.69** 25.68** 18.57* 7.45 9.81** 32.03* MS 3783 x DA 94-6 1.03 -1.97 22.88** 3.44 36.66** 7.33* 26.64** KPMS 1050 x DA 93-2 -2.86 -3.81 7.48 38.40** 58.74** 9.52** 25.78** DAMS-1 x DA 37 -0.61 12.56** -0.51 54.59** 4.40 7.79** 25.53** MS 3783 x DA 93-1 -1.52 -0.64 1.44 3.79 44.99** 7.12** 16.64* SE ± 2.00 2.76 9.43 9.69 10.33 0.96 8.88 *Significant at P = 0.05. **Significant at P = 0.01 Table 15. Specific combining ability effects in the crosses for seed yield and component traits in long duration pigeonpea Pusa Pandey (2004) 55

Slide 56: 

Sources of variation df Seed yield(g/pl) 100 seed weight(g) 50% flowering 75% maturity (days) Plant height(cm) Primary branches per plant Pods per plant F 1 generation GCA 10 506.02** 1.809** 182.755** 121.336** 2080.48** 32.934** 19058.1** SCA 55 218.75** 0.771 61.322** 41.145** 341.11** 19.713** 5934.17** Error 65 39.17 0.655 1.821 18.239 59.88 0.747 814.86 Б 2 g 44.20 0.16 18.68 12.34 267.60 2.03 2019.07 Б 2 s 179.57 0.12 59.50 22.91 281.24 18.97 5119.31 PR 0.49 2.74 0.63 1.08 1.90 0.21 0.79 F 2 generation GCA 10 242.15** 0.59 111.28** 30.78 812.73** 8.84** 2246.88** SCA 55 130.84** 0.68 51.89** 21.04 421.36** 2.87** 1179.87** Error 65 83.10 0.47 24.01 70.21 36.71 0.61 13.72 Б 2 g 17.13 0.72 9.14 1.50 60.21 0.92 164.16 Б 2 s 47.73 0.20 27.88 -49.17 384.65 2.26 1166.15 PR 0.72 -0.13 0.66 -0.06 0.31 0.81 0.28 *,** Significant at 5 and 1 percent level respectively Table 16. Analysis of variance for combining ability for yield and its attributes in F1 and F2 generation of pigeon pea Khargone. Kandalkar(2005) 56

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Genotypes Seed Yield g/plant 100 seed weight 50 % flowering Plant height Primary branches per plant Pods per plant GAUT 9002 2.63 0.66* 0.71* -12.85** 1.15** 1.24 Pant 142 -7.83** 0.35 -2.52** -16.85** -1.75** -50.99** GAUT 8630 8.20** -0.55* 1.33** 18.61** 2.90** 4.55 JJAL 16 -2.15 0.34 6.33** 5.76** 0.49* 4.48 KE 1 -2.56 0.381 -5.13** 6.61** -1.62** 50.55** PBNA 67-1 -5.42** -0.22 1.25** -12.31** -0.21 -69.91** UPAS 120 -6.35** -0.31 -4.29** -11.93** -1.93** -18.37* ICPL 87119 11.07** -0.15 1.64** 10.38** 1.25** 35.71** KM 84 3.97* -0.05 3.87** 9.76** 1.56** 33.78** JKM 7 3.31* -0.24 1.79** 12.07** -0.57* 34.32** PUSA 942 -4.87 -0.19 -4.98** -9.24** -1.26** 25.37** SE (gi) 1.66 0.21 0.36 2.05 0.23 7.55 *,** Significant at 5 and 1 percent level respectively Table 17. Estimates of general combining ability effects in F 1 generation of pigeon pea for yield and its attributes Khargone. Kandalkar(2005) 57

Slide 58: 

Promising crosses Seed Yield g/plant 50 % flowering Plant height Primary branches per plant Pods per plant GAUT 9002 x GAUT 8630 37.92** -5.60** -11.51 -1.86* 9.48* GAUT 8630 x PUSA 912 30.92** 2.09 52.87** 18.54** 89.67* GAUT 8630 x ICPL 87119 27.78** -6.53** -11.74 -1.96* 78.59* KM 84 x PUSA 942 25.15** 0.55 3.72 0.89 190.44** GAUT 9002 x ICPL 87119 22.56** 3.09* 4.72 1.79* 41.90 Pant 142 x ICPL 87119 22.52* 1.32 11.72 3.69** -25.87 GAUT 9002 x KE 1 21.68** -4.14** -19.51* -2.64* -52.95 GAUT 9002 x JJAL 16 21.28** 5.40** 24.33* -2.46* 78.13* KE 1 x PUSA 942 15.18* 0.55 -0.13 -2.93* -56.33* GAUT 8630 x JKM 7 14.24* 21.32** 16.56* -3.75** -105.03 SE(S ij ) 6.03 1.30 7.46 0.83 27.52 *,** Significant at 5 and 1 percent level respectively Table 18. Estimates of specific combining ability effects in F 1 generation of pigeon pea for yield and its attributes Khargone. Kandalkar(2005) 58

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Source of variation df Days to 50% flowering Days to maturity Plant height (cm) No. of primary branches/ plant No. of clusters / plant No. of pods / plant 100 seed weight (g) Seed yield plant (g) Replications 2 6.39 2.61 46.39 0.66 18.02 610.53 0.35 14.29 Treatments 33 372.69** 373.08** 403.50** 13.50** 403.21** 6157.64** 5.73** 99.22** Parents 9 499.55** 522.09** 652.73** 12.64** 412.48** 5837.42** 9.47** 79.98** Parents vs. Crosses 1 634.09** 453.96** 1815.07** 56.74** 15.66 11835.38** 3.32** 226.97** Crosses 23 311.68** 311.26** 244.61 11.96** 416.44** 6036.12** 4.38** 101.19** Lines 5 551.06** 837.14** 367.28 11.70 447.21 9226.47 11.95** 63.79 Testers 3 1010.28** 514.98* 541.38* 38.83** 644.99 9420.85 7.67** 385.40** Lines x Testers 15 92.16** 95.23** 144.38 6.67** 360.47** 4295.74** 1.19** 56.82* Error 66 3.42 2.71 194.44 1.31 28.10 1586.31 0.22 25.61 **Significant at 1% level, * Significant at 5% level Table 19. Analysis of variance for combining ability of the characters studied in Line x Tester analysis in pigeon pea Tandur Kumar et al . (2009) 59

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Days to 50% flowering Days to maturity No. of primary branches/plant No. of clusters/ plant No. of pods /plant 100 seed Weight (g) Seed yield /Plant (g) Lines PRG-100 -0.22 -2.76** 1.60** 6.05** 33.79** 1.27** 3.48* PRG-88 -0.06 -5.68** -0.57 -7.64** -1.88 0.74** -2.15 LRG-30 10.03** 15.99** 0.76* 6.23** 32.62** 0.20 3.68* LRG-38 -1.89** -1.68** -0.87* -3.20* -41.20** -0.90** -2.13 ICPL-85034 -10.81** -7.01** -0.80* -5.27** -22.48 -1.40** -2.87 ICPL-85063 2.94** 1.15* -0.12 3.83* -0.85 0.10 -0.02 SE (gi) 0.53 0.47 0.33 1.53 11.49 0.14 1.46 Testers ICP-8863 -3.36** -1.04** 0.36 -0.38 20.41* 0.54** 4.28** ICP-84036 -1.03* -3.38** -1.09** -3.65** -20.67* -0.62** -5.32** ICP-87119 10.75** 7.85** 1.93** 8.57** 19.19* 0.59** 3.44** ICP-89044 -6.36** -3.43** -1.19** -4.54** -18.93* -0.51** -2.40* SE (gj) 0.44 0.39 0.27 1.25 9.39 0.11 1.19 ** Significant at 1 per cent level, * Significant at 5 per cent level Table 20. Estimates of general combining ability effects for lines and testers in pigeon pea Tandur Kumar et al . (2009) 60

Slide 61: 

Cross Days to 50 % Flowering Days to maturity No. of primary branches per plant No. of clusters per plant No. of pods per plant 100 Seed weight Seed yield per plant 1 LRG-30 x ICP-87119 -2.25** 1.07 0.90 4.48 52.52* 0.64* 6.96* 2 LRG-30 x ICP-8863 4.53** -0.38 2.16** 14.79** 49.54* 0.63* 6.77* 3 PRG-100 x ICP-87119 3.67** 0.82 0.79 7.80* 48.35* 0.65* 6.76* 4 PRG-100 x ICP-8863 1.78 4.71** 1.36* 13.54** 47.10* 0.65* 6.74* 5 ICPL-85063 x ICP-87119 -3.83** 6.90** -1.56* -2.95 47.19* 0.64* 6.72* 6 ICPL-85063 x ICP-89044 3.28** 3.18** 1.69* 10.36** 6.98 0.72* 3.42 7 PRG-88 x ICP-84036 0.28 0.29 1.11 7.90* -10.41 0.12 2.79 8 LRG-38 x ICP 89044 0.11 -0.99 0.52 8.65** 47.19* -0.17 2.10 9 PRG-88 x ICP-87119 -1.17 -7.60** 0.62 -4.12 20.92 -0.08 0.86 10 ICPL-85034 x ICP 8863 -8.31** -6.38** -0.38 0.19 -11.29 -0.29 0.46 SE ( Sij ) 1.07 0.95 0.66 3-:06 22.99 0.27 2.92 ** Significant at 1 per cent level, * Significant at 5 per cent level Table 21. Estimates of specific combining ability effects in pigeon pea Tandur Kumar et al . (2009) 61

Slide 62: 

Source d.f. Days to 50% Flowering Days to maturity Plant height (cm) No. of branches per plant Number of pods per plant GCA 7 348.11** 828.56** 21.81.50** 45.12** 20699.24** SCA 28 145.68** 581.18** 2917.82** 10.66** 15091.71** Environments 2 7.15 77.29** 376.80** 16.26** 1894.30** GCA x E 14 14.84** 105.44** 302.90** 0.96** 394.95** SCA x E 56 18.86** 89.00** 330.70** 1.28** 237.86** Error 210 2.43 4.04 52.08 0.07 31.01 б 2 GCA 11.52 27.48 70.98 1.50 688.94 б 2 SCA 47.78 192.38 955.24 3.53 5020.234 б 2 gca / б 2 sca 0.24 0.14 0.07 0.42 0.13 *Significant at 5% level: ** Significant at 1% level Table 22. Analysis of variance for combining ability under individual environments and pooled over environments for different characters . Navsari Vaghela et al. (2009) 62

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Table 22Cont….. . Source d.f. Number of seeds per pod Pod length (cm) 100-seed weight (g) Seed yield per plant (g) Protein content (%) GCA 7 1.79** 4.32** 14.41** 2654.14** 17.06** SCA 28 1.58** 2.18** 4.84** 983.57** 22.95** Environments 2 0.02 1.80** 3.67** 697.85** 4.25** GCA x E 14 0.14** 0.50** 0.44** 23.39* 3.23** SCA x E 56 0.21** 0.40** 0.72** 42.20** 3.98** Error 210 0.02 0.04 0.09 11.29 0.49 б 2 GCA 0.05 0.14 0.47 88.09 0.55 б 2 SCA 0.52 0.'71 1.58 324.09 7.48 б 2 GCA / б 2 SCA 0.11 0.19 0.30 0.27 0.07 *Significant at 5% level: ** Significant at 1% level Navsari Vaghela et al. (2009) 63

Slide 64: 

Lines Day to 50% flowering Days to maturity Plant height (cm) Branches /plant Pods /plant Seeds/ Pod Pod length (cm) 100 seed weight (g) Seed yield /plant (g) Protein content (%) ICPL-87 - 6.46** -6.05** -1.99 0.35** 10.68** 0.11** 0.16** 0.13* 7.36** -1.26** 1CPL-88039 -3.05** -7.91** 0.94 -0.43** -0.79 0.08** 0.08* 0.03 -0.72 0.77* BSMR-736 1.36** 2.68** -8.67** 0.60** -10.03** -0.29** -0.17** -0.06 -1.02 0.62** GT-101 0.13 -0.15 0.67 0.88** -18.64** 0.09** -0.12** -.0.66** -9.35** -0.29* BDN-2 3.45** 8.06** -6.97** -1.36** 7.30* 0.14** -0.14* -0.40** -6.11** 0.09 ICPL-87119 4.00** 4.34** 9.09** 0.94** 25.89** 0.16** 0.48* 0.33** 10.72** 0.48** BANAS 0.48 0.61 -8.21** -2.19** -48.67** -0.47** -0.70** -0.78** -13.15** 0.49** GT-1 0.07 -1.58** 15.15** 1.21** 34.26** 0.16** 0.42** 1.41** 12.27* -0.91** *, ** Significant at 5 and 1 per cent level Table 23. Estimates of gca effects of parents pooled over environments for different traits in pigeon pea Navsari Vaghela et al. (2009) 64

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Hybrids SCA effects Mean seed yield/plant (g) GCA status Significant response in other traits for sca effects ICPL-88039 x BDN-2 29.24** 8.3.71 A x P DF, DM, PP, SP, TW , PC ICPL-87 x BSMR-736 26.32** 93.98 G x A BP, PP . SP, PL, TW , PC GT-101 x ICPL-87119 23.32"* 86.33 P x G PH, BP, PP , SP, PL, TW Banas x GT-1 22.35** 82.78 P x G DF, DM, PP , SP, PL, TW ICPL-88039 x ICPL-87119 20.65** 91.97 A x G BP, PP , PL, TW BSMR-736 x ICPL-87119 20.07** 91.08 A x G DF,DM, PP, SP, PL, TW , PC GT-101 x GT-1 18.83** 83.07 P x G PH, BP, PP , SP, PL, TW , PC BDN-2 x ICPL-87119 16.76' 83.68 P x G PH, BP, PP , SP, PL, TW , PC ICPL-87 x BDN-2 15.82** 73.38 G x P BP, PP , PL, TW ICPL-88039 x GT-1 11.75** 84.61 A x G PH, PP SP PL, TW ** Significant at 1 per cent level Table 24. Ten best specific combiners for seed yield/plant and their performance for other traits in pigeon pea DF= Days to 50% flowering SP= Seeds per pod BP= Branches per plant DM= Days to maturity PL= Plant length (cm) PC= Protein content (%) PH= Plant height TW= Test weight (g) PP= Pods per plant Navsari Vaghela et al. (2009) 65

GENE ACTION: 

GENE ACTION

Slide 67: 

Gene action refers to the behaviour or mode of expression of genes in a genetic population. Gene action is measured in terms of component of genetic variances or combining ability variances and effects. It is of two types viz. additive gene action and non-additive gene action. Gene action can be studied with the help of various biometrical techniques. 67

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Cross Gene effects Χ 2 Value M d h i j l Plant height UPAS 120 x ICPL 84023 160.64** 29.48** 11.87 - -32.18* - 2.42 ICPL 84023 x KPBR 80-2-1 180.83** -71.57** 126.01* - 67.99* -107.72** 2.01 ICPL 87119 x KPBR 80-2-1 242.51** -21.41** - - - - 2.95 ICPL 90035 x KPBR 80-2-1 233.48** -31.07** -8.61** - 56.78* - 0.36 Primary branches per plant UPAS 120 x ICPL 84023 10.24** 1.31* - - -10.23** - 3.42 ICPL 84023 x KPBR 80-2-1 15.70** -6.68** 5.09** - - - 7.48 ICPL87119 x KPI3R 80-2-1 21.12** -2.90 - - - - 3.89 ICPL 90035 x KPI3R 80 21.51** -5.26* - - - - 3.27 Table 25. Estimation of gene effects in four pigeonpea crosses Pantnagar Gupta et al. (1996) 68

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Gene Components Grain yield/ plant No. of effective clusters/ plant No. of pods/ plant No. of seeds/ plant 100 grain weight (g) D 134.10**± 12.66 719.50**± 86.20 4811.00**± 561.74 0.02 ± 0.02 5.86**± 0.07 H 1 1066.00**± 121.95 3104.00± 832.04 20470.00**± 5409.49 0.62*± 0.27 2.73**± 0.73 H 2 723.40**± 107.45 2577.00**± 733.15 13350.00**± 4766.52 0.47 ± 0.24 1.71** ± 0.64 F 141.90*± 60.76 153.20 ± 414.55 3850.00± 2695.20 0.03 ± 0.13 0.70 ± 0.36 h 2 203.36**± 18.04 906.88**± 123.10 3879.66**± 800.35 0.01 ± 0.03 0.12 ± 11 E 1.84 ± 4.47 2.42 ± 30.34 1.82 ± 198.60 0.02 ± 0.02 0.09 ± 0.05 (H 1 /d) 1/2 2.81 2.07 2.06 6.05 0.69 H 2 / 4 H 1 0.16 0.20 0.16 0.18 0.15 KD/ KR 1.46 1.10 1.48 1.32 1.20 H 2 / H 2 0.28 0.35 0.29 0.02 0.06 Heritability 12.59 19.55 22.44 2.47 70.37 t 0.12 0.43 1.52 3.80 0.14 69 Table 26. Estimates of genetic components of variation for grain yield and its components Badnapur Jahagirdar and Shinde (1996)

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Cross Gene effects χ 2 Value M d h i j l 100 seed weight UPAS 120 x ICPL 84023 8.22** -0.31** - - - - 8.78 ICPL 84023 x KPBR 80-2-1 7.46** 0.59** - - -3.19** - 1.37 ICPL 87119 x KPBR 80-2-1 8.03** 1.34** - - - 0.91* 6.88 ICPL 90035 x KPBR 80-2-1 7.09** 0.47** - - - - 6.52 Harvest index UPAS 120 x ICPL 84023 21.71** 0.92** -13.95** - - 15.30** 4.20 ICPL 84023 x KPBR 80-2-1 17.92** 6.24** - - 10.81** 5.52* 3.11 ICPL87119 x KPI3R 80-2-1 12.35** 3.97** 6.94** 4.51** - - 4.99 ICPL 90035 x KPI3R 80 7.52** - 8.55** 4.91** 3.40** - 3.57 *P ≤0.05 and**P≤0.01 Table 27 Estimate of gene effects for seed size and harvest index in four intervarietal crosses of pigeonpea Pantnagar Gupta et al. (1997) 70

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Components Year Days to 50% flowering Days to maturity Plant height (cm) Branches/ plant Pods/ plant Seeds yield/ plant (g) 100-seed weight (g) m I 92.40** 152.40** 189.10** 10.42** 161.70** 40.00** 8.72** II 91.95** 161.09** 134.14** 15.30** 158.06** 35.43** 9.82** d I 89.18** 149.50** 173.00** 9.51** 143.20** 33.02** 7.98** II 91.73** 151.95** 103.92** -16.61** 129.31** 41.31** 7.83** h I -191,11** -160.56** -180.63** -18.49** -189.99** -107.71** -18.96** II 176.57** -131 01** -113.52** -20.23** -65.32** -17.90** -24.89** i I -191.26** -160.60** -190.40** -22 66** -190.40** -93.96** -18.92** II -178.34** -134.46** -118.72** -27.98** -173.62** -59.10** -23.90** l I 195.66** 185.13** 198.46** 60.94** 213.99** 166.62** 29.36** 11 182.94** 147.26** 161.28** 63.29** 182.00** 152.48** 38.78** Type of epistasis I D D D D D D D II D D D D D D D *Significant at 5% level: ** Significant at 1% level; I first year. II second year. Table 28. Estimates of genetic parameters based on five generation means for seven characters in the cross MS Prahhat (DT) x Manak by Hayman (1958) model Hisar Hooda et al. (2003) 71

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Components Year Days to 50% flowering Days to maturity Plant height (cm) Branches/ plant Pods/ plant Seeds yield/ plant (g) 100-seed weight (g) m I 90.73** 143.73** 175.84** 7.93** 114.57** 37.90** 9.84** II 86.85** 159.60** 143.60** 12.62** 159.73** 35.49** 8.26** d I 88.79** 144.29** 193.78** 15.36** 188.90** 21.19** 8.40** II 90.52** 157.17** 139.85** 12.42** 163.16** 27.827 8.09** h I -185.60** -188.80** -211.88** -2.18 -102.65** -99.01** -22.62** II -158.51** -197.31** -181.50** -23.01** -212.40** -70.44** -16.69** i I -185.44** -186.34** -216.00** 100.00** -59.52** -109.22** -22.56** II -166.36** -224.06** -194.70** -25.64** -212.60** -86.32** -16.86** l I 197.09** 193.00** 251.00** 16.76** 29.21** 154.82** 35.72** II 198.02** 287.82** 196.20** 47.38** 309.88** 132.24** 34.02** Type of epistasis I D D D - D D D II D D D D D D D *Significant at 5% level: ** Significant at 1% level; I first year. II second year. Table 29. Estimates of genetic parameters based on five generation means for seven characters in the cross QMS-1 x Manak by Hayman (1958) model Hisar Hooda et al. (2003) 72

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Cross Scales Type of interaction (m) (d) (h) ( i ) (j) (l) Primary branches per plant UPAS-120 X C.scarabaeoides 27.43 8.73** -33.85 -9.89 -18.10 19.46 D PA-134 X C.scarabaeoides 30.78* 6.98** -21.68 -15.33 -3.69 2.50 D ICPL-84023 X C.scarabaeoides 14.63 1.63 12.5 -4.87 10.27 15.40 C Biological yield per plant UPAS-120 X C.scarabaeoides 655.03 154.38** -1240.44 -364.60 -189.37 905.58* D PA-134 X C.scarabaeoides 508.34* 86.15** -893.49* -253.69 -79.80 661.89* D 100- seed weight UPAS-120 X C.scarabaeoides 4.47* 3.07** -4.03 0.66 -1.79 2.60 D PA-134 X C.scarabaeoides 4.65 3.15** -5.72 0.27 -3.23* 5.23 D ICPL-84023 X C.scarabaeoides 9.78 3.25** -15.90 -4.79 -2.89 9.49* D Seed yield/plant UPAS-120 X C.scarabaeoides 40.27** 14.24* -84.60** -20.19* -22.94* 56.52** D PA-134 X C.scarabaeoides 18.39 7.89 -33.82 -1.27 10.19 32.31 D P=0.05 and 0.01 respecttively ; D= Duplicate, C= Complementary Pantnagar Singh and Bajpai (2005) Table 30. Estimate of gene effects for yield and yield attributing traits in interspecific crosses of pigeonpea 73

CONCLUSION: 

CONCLUSION 74 Magnitude of heterosis effects was varied from characters to characters and also for cross to cross for specific traits. High heterosis for seed yield and its component attributes lead to feasibility and development of hybrid varieties in pigeonpea using both GMS and CGMS. Combining ability analysis provides information about General and specific combining ability effects of parents and hybrid, respectively. Identification of suitable parents for hybridization programme and suitable crosses for heterosis breeding in pigeonpea. Nature and magnitude of gene action involved in the inheritance of seed yield and its attributes in pigeonpea.

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