Seminar on Heterosis & Combining Ability in Soya By CHANDEL

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j:

j Wel Come to Seminar Series 2012

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2 HETEROSIS AND COMBINING ABILITY IN SOYBEAN Seminar on

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3 HETEROSIS AND COMBINING ABILITY IN SOYBEAN MAJOR GUIDE Dr. N. B. PATEL Assistant Professor Department of Genetics and Plant Breeding Junagadh Agricultural University Junagadh 362 001 (GUJARAT) SPEAKER Kamlesh Kumar Chandel M.Sc. (Agri.)

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 4 INTRODUCTION HETEROSIS AND ITS IMPORTANCE COMBINING ABILITY AND ITS IMPORTANCE ACHIEVEMENT CONCLUSION CONTENTS

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TAXONOMY Common name : Soybean Botanical Name : Glycine max L. (1737) Family : Fabaceae Subfamily : Papilionaceae Chromosome Number : 2n=40 HETEROSIS AND COMBINING ABILITY IN SOYBEAN 5

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Origin of Soybean CHINA JAPAN KOREA RUSSIA MONGOLIA Manchuria Soybeans came from Manchuria , A region of north china HETEROSIS AND COMBINING ABILITY IN SOYBEAN 6

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INTRODUCTION HETEROSIS AND COMBINING ABILITY IN SOYBEAN 7 Soybean ( Glycine max L. Merrill) is considered a miracle crop because of its dual qualities, viz., high protein and oil content in seed. In India, soybean cultivation was introduced in 1977. It has high yield potential, wide adaptability, short duration and very high nutritional value having a vast multiplicity of uses as food and industrial products. Being a legume, it fixes a large amount of atmospheric nitrogen in soil. Therefore, soybean crop is known as “ Golden Bean ”, “ Miracle Crop ”, “ Wonder Crop ” and “ Gold of Soil ” etc.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 8 Soybean contains 40-44 per cent protein and 20-23 per cent edible oil and 30-35 per cent carbohydrate. Soybean crop very small proportion is consumed directly by humans. Raw soybean 1 , including the immature green form, are toxic to humans, swine, chickens, in fact all mono-gastric animals. Cont.... 1. Raffinoe and trypsin inhibitors

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 9 Dolichos soja L. Glycine angustifolia Miq . Glycine gracilis Skvortsov Glycine hispida ( Moench ) Maxim. Glycine soja sensu auct . Phaseolus max L. Soja angustifolia Miq . Soja hispida Moench Soja japonica Savi Soja max ( L. ) Piper Soja soja H.Karst . Soja viridis Savi Wild relatives

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 10 World Production of Soybean Soybean Producers in 2011 (Million metric tons) 1 United States 83.2 2 Brazil 72.0 3 Argentina 48.0 4 China 13.5 5 India 11.0 5 Paraguay 6.4 7 Canada 4.2 8 Others 13.1 World Total 251.5 The American Soybean Association

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 11 MAJOR PRODUCTION STATES OF INDIA Madhya Pradesh Rajasthan Chhattisgarh Gujarat Maharashtra Karnataka Uttar Pradesh Wikipedia.com

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( SOPA, 2011 ) India AREA 9.300 million hectares PRODUCTION 116.50 million tones PRODUCTIVITY 1089 kg/ha Gujarat AREA, PRODUCTION AND PRODUCTIVITY HETEROSIS AND COMBINING ABILITY IN SOYBEAN 12 AREA 8,400 hectares PRODUCTION 68,000 tones PRODUCTIVITY 810 kg/ha

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HETEROSIS HETEROSIS AND COMBINING ABILITY IN SOYBEAN 13

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HETEROSIS G. H. Shull - 1914 HETEROSIS AND COMBINING ABILITY IN SOYBEAN 14 The term heterosis was first used by Shull in 1914 . Heterosis refers to the superiority of F 1 hybrid in one or more characters over its parents. The term hybrid vigour is used as synonym for heterosis. Heterosis differs from luxuriance.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN HETEROSIS 15 For most of the characters, the desirable heterosis is positive but for some characters negative heterosis is also important in soybean. for example, Negative heterosis Days to flowering Days to maturity Plant height Toxic substance Positive heterosis Primary branches per plant Clusters per plant Pods per plant Pods per cluster Seeds per pod Seed yield per plant 100 seed weight Protein content Oil content

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 16 Superiority over parents Confined to F 1 generation Genetic control Associate with SCA Effect of heterozygosity Differs from luxuriance Low frequency FEATURES OF HETEROSIS

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TYPES OF HETEROSIS GENETIC BASIS OF HETEROSIS - Dominance Hypothesis - Over Dominance Hypothesis - Epistasis gene action PHYSIOLOGICAL BASIS OF HETEROSIS MOLECULAR BASIS OF HETEROSIS 17

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 18 GENETIC BASIS OF HETEROSIS Dominance Hypothesis Proposed by Davenport (1908) At each locus the dominant allele has a favorable effect and recessive allele has an unfavorable effect. In heterozygous state, the deleterious effects of recessive alleles are masked by their dominant alleles. Heterosis results from the masking of harmful effects of recessive alleles by their dominant alleles.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 19 Cont.... Heterosis is the result prevention of expression of harmful recessives by their dominant alleles. Similarly, inbreeding depression is a result of the homozygosity of recessive alleles, which have harmful effects. It should be possible to isolate such inbreds that have all the dominant alleles present in the population. Such inbreds would be as vigorous as the open-pollinated varieties.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 20 1. Failure in the Isolation of Inbreds as Hybrids Isolation of inbreds with all the dominant genes have not been isolated in many studies. 2. Asymmetrical Distribution in F 2 In F 2 , dominant and recessive characters segregate in the ratio of 3:1. therefore, should not show a symmetrical distribution in F 2 . OBJECTIONS

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 21 Overdominance Hypothesis This hypothesis was independently proposed by East and Shull (1908) and the term was coined by Hill (1945). This is sometimes also known as Single gene heterosis Super dominance Cumulative action of divergent alleles Stimulation of divergent alleles.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 22 Heterozygotes at some of the loci are superior to both the relevant homozygotes. Heterozygotes for more divergent alleles would be more heterotic. The heterozygote over both homozygotes may arise either due to - Production of superior hybrid substance in heterozygote which is completely different from either of the homozygous products Cont....

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 23 Cont.... - Greater buffering capacity in the heterozygote resulting from cumulative action of divergent alleles or stimulation of divergent alleles. Evidence for Overdominance In case of maize, gene ma affects MATURITY . The heterozygote Mama is more vigorous than the homozygotes MaMa and mama . EPISTASIS GENE ACTION Interaction between alleles of two or more different loci.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 24 Related with Embryo and seed sizes of species . Growth rates in the various stages of development of species . Rates of reproduction of species Rates of various assimilation activities of species . PHYSIOLOGICAL BASIS OF HETEROSIS

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 25 Some reports of heterosis for physiological character FINDING AUTHOR (S) Explain heterosis in maize for hormone balance, crop contain higher amount of GAs than their inbred parents. Rood et al. (1988, 1990) Heterotic hybrids generally show a faster growth rate, higher leaf area index and greater biomass production than their parents. Chopra et al. (1993) Photosynthetic efficiency in rice due to positive N 2 content. Ankita et al . (1988)

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 26 MOLECULAR BASIS OF HETEROSIS MECHANISMS Intermediate amount of a single gene product Such situation occurs when the homozygote for dominant allele produces more and the heterozygote for the recessive allele produces less than the optimum level of a product. In the bread mould Neurospora crassa . Gene pab of this mould is responsible for the production of p - aminobenzoic acid. The heterozygote pab + pab produces intermediate amounts of p - aminobenzoic acid and is faster in growth than the two homozygotes, that is, pab + pab + and pab pab .

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 27 Separate Gene Products Two alleles of a gene may produce proteins with slightly different functions and properties. The heterozygote for such a gene would produce the proteins encoded by both the alleles. Sickle cell anemia in human beings . The homozygote ss effected person does not survive in life. The heterozygote Ss is more resistant to malaria than the normal homozygote SS and ss. Cont....

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 28 This is because the defective haemoglobin β s produced by the recessive allele s is not suitable for the multiplication of malarial parasite, which multiplies in the red blood cells. Thus the heterozygotes survive due to the normal haemoglobin p A produced by the normal allele S, and are resistant to the malarial parasite due to the defective haemoglobin p s produced by the recessive allele. Cont....

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FIXATION OF HETEROSIS HETEROSIS AND COMBINING ABILITY IN SOYBEAN 29 Asexual Reproduction Heterosis can be easily conserved in Vegetatively Propagated Crops Such as Sugarcane, Potato, Sweet Potato, Banana etc. Apomixis The fixation of heterosis by apomixis is common in Citrus fruits, blackberries, roses, bluegrasses and many other flowering plants. As a result of apomixis, Mandal could not confirm his result findings of garden peas on Hieracium spp. . The F 2 and later generation progenies were similar to their F 1 hybrid. Hieracium spp.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 30 Balanced Lethal System BLS leads to fixation of heterosis in some plants. In Oenothera spp. Genetic segregation is almost completely suppressed by BLS. The homozygote ss are lethal, hence they die. Only hetrozygoutes Ss survive. Polyploidy – Chromosome Doubling Heterosis can also be fixed by chromosome doubling or polyploidy. Especially in interspecific and intergeneric hybrids. Wheat X Rye = Triticale Cont....

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FACTORS AFFECTING HETEROSIS HETEROSIS AND COMBINING ABILITY IN SOYBEAN 31 Many genetic factors which affect magnitude of heterosis in crop plants Mode Pollination The level of heterosis is generally higher in cross pollinated species than in self pollinated species. Genetic Diversity of Parents The expression of heterosis is influenced by genetic diversity of parents. Heterosis increased with genetic divergence in morphological characters and also with respect to geographical origin of parents. Greater heterosis is associated with greater parental diversity.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 32 Genetic Base of Parents The manifestation of heterosis is affected by the genetic base of the parents. For example, in cross pollinated crops higher heterosis is associated with broad genetic base of the parents. Adaptability of Parents The magnitude of heterosis is also affected by the adoptability of the parents. In cross pollinated crops, heterosis is associated with wide adaptability of the parents, because there is closely association between adaptability and genetic base of the parents. Cont....

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN Traits that are low in heritability usually exhibit the more heterosis and the traits with higher heritability usually show little heterosis. The heritability of the trait is inversely related to how easily it is hanged through selection of parents. Traits low in heritability are very hard to improve through selection, but a lot of genetic improvement can be made in these traits through heterosis. Traits high in heritability do not show much heterosis, but are easy to change through selection. 33 LOW/HIGH HERITABILITY INTERACTION

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN ADVANTAGES 34 Heterosis (Cross breeding) offers some important advantages It has been well documented that cross breeding improves the performance of line. Virtually all commercial lines producers utilize some form of cross breeding in their genera. Like any other management technique, cross breeding must be done properly for the full benefits to be realized.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN OTHER ADVANTAGES 35 Heterosis leads to increase in yield Reproductive ability Adaptability Disease and insect resistance General vigor Quality improvement

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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 HETEROSIS AND COMBINING ABILITY IN SOYBEAN 36

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2) HETROBELTIOSIS When the heterosis is estimated over the superior or better parent, it is referred to as heterobeltiosis. It is worked out as follows Heterobeltiosis = Where , BP = mean value (over replication) of the better parents of the particular cross HETEROSIS AND COMBINING ABILITY IN SOYBEAN 37 Cont....

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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 HETEROSIS AND COMBINING ABILITY IN SOYBEAN 38 Cont....

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Case Study HETEROSIS AND COMBINING ABILITY IN SOYBEAN 39

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Estimation of heterosis in soybean over mid parent (MP) and better parent (BP) in per cents Table no. 1 Dharwad Ramana and Satanarayana (2006) * and ** Significant at 5% and 1% levels, respectively Characters Rang of heterosis Average heterosis Top heterotic crosses MP BP MP BP MP BP Harvest index (%) -12.88 to 23.12 -22.68 to 21.00 2.5 -2.58 LSb 1XPK 472(23.12**) LSb 1XPK 472(21.00**) LSb 3XPK 472(14.87**) LSb 3XPK 472(14.82**) DSb 1XPK 1029(14.51**) DSb 1XPK 1029(14.09**) LSb 3XPK 1029(13.19**) LSb 3XPK 1029(10.24*) Seed protein content (%) -5.31 to 4.22 -7.19 to 3.48 0.75 -0.59 JS90-41XPK 472(4.22) JS90-41XPK 472(3.48) JS90-41XJS 335(3.86) LSb 3X MACS 1029(2.87) LSb 3XPK 1029(3.68) JS90-41XMACS 101(2.80) JS 90-41XMACS 201(3.40) DSb 1XPK 472(1.62) Oil content (%) -6.77 to 4.94 -10.80 to 4.10 0.63 -1.54 LSb 3X MACS 201(4.94*) LSb 3X MACS 201(4.10) LSb 3XPK 472(4.94*) JS90-41XMACS 201(3.50) JS90-41XMACS 201(4.11) LSb 3XPK 472(3.04) LSb 3XPK 1029(3.68) - Grain yield per plant (g) -18.08 to 43.41 -39.34 to 34.69 16.85 1.22 DSb 1XPK 1029(43.41**) DSb 3XPK1029(34.69**) JS90-41XPK 472(39.41**) LSb 3XPK 472(32.90**) DSb 1XPK 472(39.08**) DSb 1XPK 472(27.48**) LSb 3XPK 472(36.45**) DSb 1XMACS 201(19.93**) The most important attribute of a plant its yielding ability. In this table cross DSb 1 x PK 1029 significant positive heterosis over mid parent and cross DSb 3 x PK 1029 significant positive heterosis over better parent for grain yield / plant. 40

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Characters Heterosis Over mid parents Over better parents Cross I ‘MACS 684 X MACS 124’ Plant height 4.85* 1.23 Pod number 2.65 1.48 Oil concentration -2.26 -4.60* Yield Kg / ha 57.97** 48.19** Cross II ‘MACS 684 X RSC 1’ Plant height -1.97 -8.70** Pod number 44.05** 31.18** Oil concentration -2.22 -2.91 Yield Kg / ha 29.18 ** 10.97** Cross III ‘PK 472 X RSC 2 Plant height 27.74** 10.03** Pod number 1.82 -15.66** Oil concentration -1.53 -7.16* Yield Kg / ha 9.27** 7.55* Heterosis (%) over mid-parent and better parent in three soybean crosses for four quantitative traits * and ** Significant at 5% and 1% levels, respectively Pune Bhosle et al. (2005) Table no. 2 The chances of getting batter selections for further generation are more in cross III as compared to the other two crosses. Because of low inbreeding depression in this cross. 41

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Characters Best heterotic crosses (%) Common crosses for SCA and heterosis (BP/EP) Better Parents Economic Parents Heat tolerance PK327 X MACS58 -29.10 ** JS80-21 X NRC21 -41.49** - JS80-21 X NRC12 -27.78** PK327 X MACS58 -36.79** - JS80-21 X PK472 -19.40** PK327 X NRC12 -33.73** - PK327 X NRC12 -18.20** JS80-21 X PK472 -31.59 ** - Germination stress index PK327 X MACS58 14.62** Bragg X MACS58 16.32** JS79-81 X PK472 NRC2 X MACS58 10.01** PK327 X MACS58 15.36** JS80-21 X PK472 Pusa20 X MACS58 9.89** JS80-21 X PK472 15.16** - JS80-21 X PK247 9.20** JS79-81 X PK472 13.55** - Crosses with significant desirable heterosis over better parent and best check variety (per cent) Udaipur Krishnaknawat and Maloo (2004) * and ** Significant at 5% and 1% levels, respectively Table no. 3 The cross PK 327 x MACS 58 and JS80-21xNRC 21 was showed significant value in desirable direction over BT, EP, and check parents for heat tolerance. Positively significant value for GST has exhibited in cross PK 327 x MACS 58 .while PK 327 x MACS 58 over BP and EP respectively. It is showed highest % of heterobeltiosis. Signif . Heterosis indicated involvement of non add. Gene action for HT(n:d) and GSI(p:d) 42

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Estimation of heterosis (%) over better parent in soybean S. No Hybrid Days to maturity No. of pods 100 seed weight (g) Seed yield (g) 1 CO 1 X MACS 124 -3.79** 12.91** -4.76** 11.16** 2 CO 1 X Bragg -4.55** -41.47** -0.70** -58.68** 3 CO 1 X EC 4296 -5.44** 20.31** 1.64** 14.91 ** 4 CO 1 X CO 2 -4.92** 5.36 -7.25** 5.75** 5 CO 1 X EC 9472 -3.03** 28.28** 1.64** 33.54 **(HXL) 6 MACS 124 X Bragg - 7.75** 6.42 -11.90** 1.11** 7 MACS 124 X EC 4296 -2.21** 2.55 -6.28 ** 3.17** 8 MACS 124 X CO 2 -5.79** -2.48 -8.74** -9.14** 9 MACS 124 X EC 9472 -2.32** -26.08** -20.13** -38.29** 10 Bragg X EC 4296 -4.43** -15.11** -6.35** -10.65** 11 Bragg X CO 2 -12.0** 30.64** -12.58** 19.39 ** 12 Bragg X EC 9472 -0.78** 7.18 -13.18** 2.61** 13 EC 4296 X CO 2 -11.07** -6.22 -7.68** 1.06** 14 EC 4296 X EC 9472 - 7.72** 14.40** -0.25 13.76 ** 15 CO 2 X EC 9472 -7.03** -2.53 -20.68** -7.79** Table no. 4 Coimbatore Ganesamurthy and Seshadri (2004) * and ** Significant at 5% and 1% levels, respectively 43 The heterotic hybrids for yield per plant of co 1 x EC 9472, Bragg x CO 2 and CO 1 x EC 4296 showed significant heterotic effects for component character number of pods, day to maturity and seed weight. Obviously, heterosis for seed yield in these crosses seems to be due to heterotic effects of component character.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 44 Combining Ability

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN The concept of combining ability as 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. COMBINING ABILITY Sprague 1942 45

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 46 Combining ability analysis helps in the evolution 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 hybrids cross combinations. For combining ability analysis, cross have to be made either in diallel or partial diallel or line X tester fashion. Combining ability analysis helps in the identification of superior hybrid combinations which may be utilized for commercial exploitation of heterosis. Combining ability analysis is a useful tool in the development of synthetic varieties. Cont....

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 47 Combining ability estimates are free from genetic assumptions and are based on empirical results ( Simmonds , 1979) Combining ability estimates of gca and sca effects are based on first order statistics. Combining ability analysis provides information about the gene action involved in the expression of various quantitative traits and thus helps in deciding the breeding procedure for genetic improvement of such traits. Cont....

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN TYPES OF COMBINING ABILITY 48 1. General combining ability ( gca ) The average performance of a stain or genotype in a series of hybrid combinations is termed as general combining ability ( gca ). The gca variance is primarily a function of the additive genetic variance, but if epistasis is present gca will also include additive X additive type of non-allelic interaction.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 49 It is estimated from half-sib families. The gca variance has positive correlation with narrow sense heritability. the gca helps in the selection of suitable parents for hybridization. Cont.... Gca

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 2. Specific combining ability ( sca ) 50 The performance of a parent in a specific cross is known as specific combining ability. sca refers to the deviation of a particular cross from the general combining ability. The sca is estimated from full sib families.

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 51 The sca variance has positive association with heterosis or hybrid vigour. The sca helps in the identification of superior cross combinations for commercial exploitation on heterosis. The sca variance is mainly a function of dominance variance, but if epistasis is present, it would also include additive X additive, additive X dominance and dominance X dominance types of non-allelic interactions. Cont.... Sca

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 52 GCA SCA Its average performance of a stain in a series of crosses It refers to the performance of specific cross in relation to gca gca is due to additive genetic variance and additive X additive epistasis sca is due to dominance genetic variance and all the three types of epistasis Its estimated from half-sib families Its estimated from full-sib families It helps in the selection of suitable parents for hybridization It helps in the identification of superior cross combinations It has relationship with narrow sense heritability It has relationship with heterosis Differences Between GCA and SCA

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Case Study HETEROSIS AND COMBINING ABILITY IN SOYBEAN 53

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Estimate of the effects of general combining ability (GCA) for the traits evaluated in the experiment involving eight parents in a diallel cross. Parents Days to flowering Plant height Number of branches Days to maturity Doko 11.539 9.779 2.275 -0.366 Bossier -6.367 -10.923 -1.315 -3.734 Ocepar – 4 -4.975 -8.520 -0.758 0.507 BR-15 2.301 -0.797 0.828 4.478 FT- Cometa -8.565 5.922 -1.119 -3.635 Sevana 6.757 10.666 0.934 0.800 Parana -4.040 -7.440 -0.897 -1.918 Critalina 7.359 6.105 0.650 5.126 E.P.( g i -g j ) 0.429 1.172 0.267 2.276 Brazil Antonio et al . (2006) Table no. 1 Parent FT – Cometa attained lowest GCA values in the sense of earliness of flowering. In relation to the traits PH, parents Bossier attained the highest negative gca values. Parents Doko aresented the highest positive gca values for the no. of branches. The lowest gca values of parent Bossier in the sense of earliness plant maturation. The magnitude of estimates indicate a greater divergence in the parents doko and FT- Cometa compared to the overall mean, so that higher heterosis value would be expected in the hybrid involving one of these parents. HETEROSIS AND COMBINING ABILITY IN SOYBEAN 54

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Crosses with significant desirable SCA effects Udaipur Krishnaknawat and Maloo (2004) * and ** Significant at 5% and 1% levels, respectively Table no. 2 The crosses JS79-81xPK472 (Good x Poor) Higher negative sca effects and considered good specific cross combination for HT. The cross JS 79-81 x PK472 (Good x Poor) was showed tolerant to moisture stress as indicated by their signif . Sca effect in desired direction. Only cross JS79-81 x PK472 was common and possessed most promising heterotic diverse parent with high sca effects showed desirable combining ability as well as high per se performance appeared for both traits . Characters Best SCA cross Status of the parents Common crosses for SCA and heterosis (BP/EP) Heat tolerance MAC124XPK427 -10.88** (PoorXPoor) - JS79-81XPK472 -9.11** ( GoorXPoor ) - BraggXMACS58 -8.41** ( GoorXPoor ) - JS71-05XPK472 -7.88** ( PoorXPoor ) - Germination stress index JS79-81XPK472 7.83** ( GoorXPoor ) JS79-81 X PK472 JS80-21XPK472 7.47** ( GoorXPoor ) - PK471XNRC12 6.97** ( PoorXAv .) - PK564XNRC12 6.41** (PoorXPoor) - 55

Analysis of variance (mean squires) for combining ability for seven characters in soybean:

Analysis of variance (mean squires) for combining ability for seven characters in soybean Source d.f . Days to flowering Days to maturity Plant height Pods per plant Grain yield per plant Protein content Oil content GCA 6 81.3** 61.4** 686.9** 2175.0** 87.2** 4.29** 3.06** SCA 21 14.7** 21.2** 91.6** 448.7** 30.3** 3.07** 1.06 Error 54 0.5 0.4 0.9 3.0 0.3 0.26 0.80 Relativ impo . 0.5 0.3 0.6 0.5 0.3 0.09 0.63 * and ** Significant at 5% and 1% levels, respectively Dharwad Gadag et al. (1999) Variance due to gca and sca ware significant for all trait except oil content, for which mean sq. due to sca was not significant for this trait. in the combining ability analysis, if both gca & sca effects are significant, it is useful to know how important the interactions are in determining the single cross progeny. Lower value of the ratio for protein content ..Predominant role of non additive gene effects in the expression of these characters. For other traits viz., oil content, .. nearly equal importance of both additive and non add. Through none of the parent was a good general combiner for all the traits. Table No. 3 56

Ranking of parents varieties of soybean based on character means and estimates of GCA:

Ranking of parents varieties of soybean based on character means and estimates of GCA Parental varieties Table no. 4 Rank Grain yield per plant (g) Protein content (%) Oil content (%) Parent Mean GCA Parent Mean GCA Parent Mean GCA 1 KHSb-2 27.6 4.72** LBS 43.54 0.60** KHSb-2 20.95 0.45 2 DS75-62 20.6 3.47** Monetta 41.68 1.02** DS75-62 20.90 0.64* 3 LBS 15.5 0.05 Bragg 41.03 -0.10 Hardee 20.85 0.24 4 Hardee 14.7 -0.88** SL-96 39.33 39.33 SL-96 20.17 0.20 5 SL-96 12.6 -2.62 Hardee 39.22 -0.08 Bragg 19.53 0.29 6 Bragg 10.8 -0.76 KHSb-2 39.09 -0.65** Monetta 19.28 -1.02** 7 Monetta 6.9 -3.89** DS75-62 38.55 -0.99 LBS 17.72 -0.50 KHSb-2 and DS75-62 exhibited high gca effect for yield and yield contributing characters. So that these are good general combiner for grain yield. Monetta has also good general combiner for protein content trait and parent DS 75-62 significant for oil content. Through none of the parent was a good general combiner for all the traits * and ** Significant at 5% and 1% levels, respectively Dharwad Gadag et al. (1999) 57

Ranking of elite crosses of soybean based on character means and estimates of SCA:

Ranking of elite crosses of soybean based on character means and estimates of SCA Top five crosses Table no. 5 * and ** Significant at 5% and 1% levels, respectively Dharwad Gadag et al. (1999) Rank Grain yield per plant (g) Protein content (%) Oil content (%) Cross F 1 Mean SCA Cross F 1 Mean SCA Cross F 1 Mean SCA 1 6X7 24.0 9.79** 5x4 44.12 3.60** 5x2 23.27 2.19** 2 4X6 27.2 9.64** 2x6 43.90 3.13** 3x6 20.42 1.05 3 1X7 33.4 9.43** 7x4 41.72 2.10** 1x7 21.08 0.98 4 3X2 29.8 7.06** 3x6 4.22 1.57** 1x4 21.35 0.85 5 7X5 18.5 5.82** 2x7 41.47 1.03** 7x2 21.03 0.74 SE ( gi ) - 0.17 - 0.16 - 0.28 SE ( sij ) - 0.49 - 0.46 - 0.80 58 5. LBS 6. DS75-62 7. KHSb-2 1. SL-96 2. Monetta 3. Bragg 4. Hardee It is interesting to note that parent LBS not showing significant gca effect for yield but gave high and significant positive sca effects for yield trait. It is not necessary that parents having higher estimates of gca effects would also give higher sca effects in cross combination.

Estimation of GCA for different characters in soybean varieties:

Estimation of GCA for different characters in soybean varieties Plant Plant height Branches Per plant Pods per plant 100 seed weight Grain Yield per plant Days to maturity Oil content Protein content PK 564 14.26* 2.46* 17.52* 3.88* 3.24* 42.37* 6.63* 15.84* PK 416 9.14* 1.29* 14.92* 3.21* 2.59* 33.79* 6.15* 8.81* PK 472 7.34* 1.44* 7.76* 2.10* 1.16* 23.75* 4.41* 5.37* SL 160 2.68* 0.85* 8.59* 0.93* 0.93* 9.98* 1.84* 3.53* SL 152 0.94* -0.21 -6.10* -0.37* -0.42* -1.05 -0.05 -0.62* DS-75-12-1 -0.05 -0.27* -6.21* -1.33* -1.05* -11.84* -2.29* -2.23* F 87-3065 -7.50* -1.28* -6.75* -1.92* -1.33* -21.33* -3.60* -7.10* F 87-3095 -11.40* -1.81* -13.37* -2.80* -2.33* -32.08* -5.67* -10.19* SL 96 -15.40* -2.17* -16.34* -3.71* -2.79* -42.99* -7.43* -13.89 S.E. (g) 0.26 0.06 0.23 0.02 0.04 0.23 0.62 0.15 CD 5% 0.50 0.11 0.45 0.04 0.08 0.45 0.12 0.29 *Significant at 1 % level Ludhiana Sharma and Phul (1994) The genotypes viz. PK 416 and PK 564 were best general combiners for most of the yield and quality traits. PK 416 popular variety in Punjab Table no. 6 59

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Specific combining ability effects of selected crosses for important traits in soybean Grain yield per plant Pods per plant Oil content Protein content F87 – 3065 X SL 96 (LXL) DS75-12-1 X SL 96 (LXL) F87 – 3065 X SL 96 (LXL) F87 – 3065 X SL 96 (LXL) F87 – 3095 X SL 96 (LXL) F87 – 3065 X SL 96 (LXL) F87 – 3095 X SL 96 (LXL) DS75-12-1 X SL 96 (LXL) DS75-12-1 X SL 96 (LXL) F87 – 3095 X SL 96 (LXL) F87-3095 X F87-3065 (LXL) F87 – 3095 X SL 96 (LXL) F87-3095 X F87-3065 (LXL) SL 152 X SL 96 (LXL) DS75-12-1 X SL 96 (LXL) F87 – 3095 X F87-3065(LXL) SL 152 X SL 160 (MXL) PK 472 X F87-3065 (MXL) SL 152 X SL 96 (MXL) DS75-12-1 X F87-3065(LXL) SL 152 X F87-3095 (MXL) PK 416 X F87-3065 (HXL) DS75-12-1 X F87-3065(LXL) SL 152 X SL 96 (MXL) PK 416 X F87-3065 (HXL) SL 160 X SL 152 (MXL) SL 152 X F87-3065 (MXL) DS75-12-1 X F87-3095(LXL) SL 160 X SL 96 (MXL) SL 160 X SL 96 (MXL) SL 160 X SL 96 (MXL) SL 160 X SL 96 (MXL) Ludhiana Sharma and Phul (1994) Table no. 7 L – Low, M – Medium and H- High general combiners. In both these crosses indigenous (SL 96) as well as exotic (F87-3065, F87-3095) strains are involved. 60

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN 61 China Sun huan et al . (2003) The team, which began its research 20 years ago, acquired in 1993 the technology to develop a cytoplasmic -nuclear male sterile line. which passed appraisal by Chinese and American experts. Using the leaf-cutting bee as a pollinator. The proportion of seeds setting on the new hybrid can reach more than 70 percent. Repeated experiments showed that a hectare of this hybrid can yield 1,000 kilograms of soybean, 20% more than usual breeds commonly grown by local farmers. World’s first hybrid soybean variety - Hybrid Dou – 1 Now,, this hybrid is commercially grown in some area of China. ACHIEVEMENT

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In india , Developed the world famous Jawahar series of soybean varieties released from JNKVV Jabalpur (MP). JS 335 – it has a wide adaptability and high yielding is popular in MP, Maharashtra, Rajasthan, Karnataka and Andhra Pradesh covering 50% area of country. JS 93-05 – It is most popular variety for a four-seed/pod culture, early maturity, resistant to root rot, tolerant to insect pests and drought conditions. It is covering in 35% area of India popular in Madhya Pradesh, Maharashtra, Rajasthan, and Karnataka States of the Country. JS 95-60 - The first extra early maturity variety released by JNKVV has made impact in the low rainfall and upland conditions. This is resistant to root rot, insect pests and drought. JS 97-52 - Recent released variety. which is high yielding, good germinability , wide adaptable, suitable for Central Zone and North Eastern Zone. It is first multiple resistant variety showing resistance to major diseases and insect pests. The addition of these varieties have been placed soybean as first ranking oilseed crop of India resulted in changing oilseed scenario in the country. Source By - Jawaharlal Nehru Krishi Vishwa Vidyalaya , Jabalpur (MP) 62 CON…

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In Gujarat In 1983 released first two soybean varieties viz., Gujarat Soybean-1 and Gujarat Soybean-2 . The last released soybean variety is Gujarat Soybean-3 . CON… 63

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN CONCLUSION From on going discussion it can be concluded that the hybrid soybean production technology is one of the best options for increase yield, oil and protein potential of soybean. Selection should be made taking yield components into consideration in segregation. Crosses could be exploited for improving quality and quantity as they had high per se performance in breeding programme . High SCA effects are considerable interest as such combinations may give desirable transgressive segregates if the additive effects of one parent and complementary effect of the other parent act in the same direction for maximum expression of the traits. It is not necessary that parents having higher GCA effects would also give the higher SCA effects. Significant heterosis indicating involvement of non-additive gene action for the traits as well also evident from significant specific combining ability mean squares. Therefore, concrete efforts should be made to develop the hybrids which can suitable for cultivation and higher productivity. 64

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HETEROSIS AND COMBINING ABILITY IN SOYBEAN THANK YOU

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