Genotype x environment interaction effects on estimations of genotypic and phenotypic parameters were studied, based on seed yield of 97 progenies and three test cultivars of common beans at two different locations in Minas Gerais, Brazil. At one location, Patos de Minas, F7 progenies and test cultivars were evaluated using a simple 10 x 10 lattice design, whereas F8 progenies and test cultivars planted at Sete Lagoas at two density rates comprised two independent experiments with the same statistical design as the former. Results indicated the need to evaluated bean progenies at several environments. This was supported by the fact that the narrow sense heritability value, estimated through covariance analysis, was similar to the realized heritability value, and the genetic expected gain estimate was practically equal to the realized gain estimate. Also, selection effectiveness based on the mean over environments, was three times greather than when based on each individual environment

ABREU, A. de F. B.

PEREIRA FILHO, I. A.

RAMALHO, M. A. P.

SANTOS, J. B. dos

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Rev. BrasiL Genet. 13, 1, 75-82 (1990)(BraziL J. Genetics)PER Fº,I.A.1990EFFECT OF GENOTYPE X ENVIRONMENT INTERACTION ONESTIMATIONS OF GENETIC AND PHENOTYPIC PARAMETERSOF COMMON BEANS *Angela de Fátima Barbosa Abreu', Magno Antonio Patto Ramalho, João Boscodos Santos2 and Israel Alexandre Pereira-FilhoêABSTRACTGenotype x environment interaction effects on estimations of genotypic and phenotypicparameters were studied, based on seed yield of 97 progenies and three test cultivars of common beansat two different locations in Minas Gerais, Brazil. At one location, Patos de Minas, F7 progenies and testcultivars were evaluated using a simple 10 x 10 lattice design, whereas F8 progenies and test cultivarsplanted at Sete Lagoas at two density rates comprised two independent experiments with the same statis-tical design as the former. Results indicated the need to evaluate bean progenies at several environments.This was supported by the fact that the narrow sense heritability value, estimated through covarianceanalysis, was similar to the realized heritability value, and the genetic expected gain estimate was practi--ally equal to the realized gain estima te. Also, selection effectiveness based on the mean over environ-+ments, was three times greater than when based on each individual environment.INTRODUCTIONIn Brazil, common beans are grown under quite diverse conditions both interms of soil and planting systems. On this basis, genotype x environment interactionbecomes a very important factor in breeding programs since the existence of interac-* Part of thesis presented by AF.B.A to Escola Superior de Agricultura de Lavras in partial fulfillmentof the requirements for the Master's degree.1 EMBRAPA/EPAMIG, Caixa Postal 176, 37200 Lavras, MG, Brasil. Send correspondence toAF.B.A.2 Departamento de Biologia, Escola Superior de Agricultura de Lavras, Caixa Postal 37, 37200 Lavras,MG, Brasil.3 EMBRAPA, CNPMS, Caixa Postal 151, 35700 Sete Lagoas, MG, Brasil.76 Abreu et al.tion implies the recommendation of cultivars for certain environments or the searchfor genetic materiaIs that are less affected by environmental variation (Santos, 1980).Interaction also affects the estimation of genetic parameters and conse- -quently the gain expected through selection. In the specific case of common beans,the information available in the literature about the major gene actions involved inthe control of seed yield is still scarce and often discordant, probably owing to thefact that interaction is not taken into consideration when estimates are calculated. Ithas been shown that when cultivars are tested under conditions of greater plant com-petition at density rates similar to those used for crop planting, additive variancethe maio component of genetic variance (Ouiâones, 1969; Hamblin and Evans, 1976;Hamblin and Morton, 1977; Santos et a/., 1985). In contrast, when cultivars are plantedat lower density rates with lower plant competition, manifestation of dominance hasbeen observed for seed yield (Chung and Stevenson, 1973; Albuquerque and Vieira,1974; Hamblin and Morton, 1977; Sarafi, 1978; Foolad and Bassiri, 1983).For common beans, major emphasis has been placed only on the identifica-tion of more stable material obtained in trials involving different cultivars (Santos,1980; Beaver et al., 1985; Pacova et a/., 1987; Park, 1987), with few studies on the ef-fect of interaction on the response to selection. Thus, the objective of the present in-vestigation was to test bean progenies at different locations, times and density ratesto quantify progeny x environment interaction and to determine its effect on the es-timations of genetic and phenotypic parameters to be used in breeding programs.MATERIAL AND METHODSThe progenies used in the present study were obtained from crosses betweenRio Tibagi and Carioca 80. The Rio Tibagi cultivar, of type 11 growth habit, is ~straight -standing plant with small black seeds and is susceptibleto anthracnose, whi,Carioca 80 is of type III growth habit, is prostrate, has cream-colored seeds with brownstripes and is resistant to anthracnose.Carioca 80 and Rio Tibagi were crossed in 1984 according to the methodproposed by Vieira (1967) and the population obtained was taken to the Fs genera-tion by the mass method. Two thousand seeds of this generation (F6) were planted,all of them having seeds of the Carioca 80 type. Of the progenies obtained, 97 show-ing straighter stand were selected and tested for seed production together with threetest cultivars (Carioca 80, Rio Tibagi and Carioca 1030) in the F7 and Fg generations.The F7 generation was tested in Patos de Minas in February 1987 using a simpIe 10 x10 lattice designo Each plot consisted of a 2-meter line planted with 15 seeds per linearmeter. Plots were spaced 0.5 m apart. The Fg generation was planted at two densityrates,8 and 16 plants per meter, at Sete Lagoas, MG, in two independent experiments,also using a simple 10 x 10 lattice designo Plots consisted of a 4-meter line and werespaced 0.5 m apart.Genotype x Environrnent Interaction in Dry Beans 77Data were analyzed by separate and joint analysis of variance consideringthe two densities and the two generations. Estimations of genetic and phenotypicvariance components were obtained according to the scheme proposed byVencovsky(1987). Selection efficiency (SE) was estimated by thefoUowing expression, proposedby Hamblin and Zimmermann (1986):SE = [(A-C)/(B-C)] . 100.re:"----'A: number of selected progenies common to the two selection generations;B: total number of progenie selected;C: number of progenies common to the two generations expected to occurstrictly at random, and considered to be 10% of B in this specific case.Realized heritability and realized gain through selection were also obtainedas suggested by Fehr (1987).RESUL TS AND DISCUSSIONIt should be pointed out that, in the present paper, the generation effectrefers to environmental factors such as planting time and location since generationswere grown at different times and locations. The generation effect itself should benonsignificant because in generations F7 and Fg the material is already practicallypure. Even in populations in the initial stages of segregation such as F2 and F3, no in-tPfaction occurs with generation (Ramalho et ai., 1988) as long as generations are--.->tedat the same planting time and location.Progeny performance was good in generation F7, with mean yields of 1812kg/ha and a range of variation of 816 to 21)87 kg/ha, thus indicating the productivepotential of some progeny and the possibility of successful selection. In generationFg, mean yield was 1708 kg/ha at both density rates, with 16% of the progenies per-forming better than the test cultivars.Joint analysis of variance for the Fg progenies at both density rates showedsignificant progeny, density and within-progeny density effects (Table I). Progeny xdensity rate interaction was not significant, showing that the material behaved similar-ly at both density rates: This absence of significant interaction was probably due tothe fact that the progenies tested had been previously selected for type of stand. In-deed, Nienhuis and Singh (1985) and Westermann and Crothers (1977) reported thatthe effect of plant density depends on the type of growth of the cultivar used, i.e., cul-tivars of determinate growth suffer less competition at higher densities than plants ofindeterminate growth.78 Abreu et ai.Table I - Joint analysis ofvariance for seed yield data (kg/ha) obtained in trials of the F8 progenies of theCarioca 80 x Rio Tibagi cross. Sete Lagoas, MG, 1987.FV d.f. MSProgenies and test cultivars (P and T)Progenies (P)Test cultivars (T)PvsT(Pand T) xDD/PDfTPxDTxD(99)962194081.5350· •93660.9075··(99)16177.6650310269.57_61953.8000391180.2250· •Effective mean error%216267963.217563262.350019105.072556810.9550P mean (kg/ha)T mean (kg/ha)CV(%)1709.501938.7113.89• and •• , F test significant at the 5 and 1% levei, respectively.Joint analysis of variance for F7and F8progenies showed signiticant progenvenvironment and progeny x environment interaction effects (Table 11).The varian.Lcomponent of progeny x environment interaction was also found to be approximate-ly double the genetic covariance between progenies at the two locations (Table 111),reflecting the true genetic variance between progenies regardless of the environmentswhere they were tested. The main implication of an interaction of this magnitude isthat it reflects directly on the response to selection. Thus, when the 20 best progenieswere selected from the average for the two sites there was coincidence of 12progeniesat Patos and of an equal number at Sete Lagoas, whereas if selection had been ap-plied at Patos, only tive progenies would have also been among the best ones testedat Sete Lagoas (Table IV). By applying Hamblin and Zimmermann's expression(1986), this corresponds to a mean selection efticiency of 55.6%, i.e., approximatelythree times that obtained if selection had been applied in one environment and gainhad been observed in another (only 16.7%). Thus, in this situation the best criterionfor progeny selection is mean progeny performance under different environmentalconditions, as proposed by Rosielle and Hamblin (1981).Genotype x Environment Interaction in Dry Beans 79Table 11- Joint analysis ofvariance for seed yield data (kg/ha) obtained in trials ofthe F7 and Fs progeniesof the Carioca 80 x Rio Tibagi cross. Patos de Minas and Sete Lagoas, MG, 1987.FV d.f. MS96 97007.5332··1 513590.0600··96 70301.71n··324 43567.62881760.9011.85Progenies (P)Generations = environments (E)Etréctive mean errorMean(kg/ha)CV(%)•• , F test significant at the 1% leveI.Table III - Estimates of genetic and phenotypic parameters for mean seed yield (kg/ha) obtained inprogeny test trials for the Carioca 80 x Rio Tibagi cross.Genetic and Phenotypic Parameters EstimatesGenetic variance among F7 progeniesGenetic variance among Fs progeniesvcÍletic covariance among Fs progenies at planting61601.320018424.9763densities of 8 and 16 plants per meterGenetic variance among progenies15199.273826719.952213352.90n26734.0889120331.095011.1017.393.154.94Genetic variance among progenies at the two locationsVariance of progeny x environment interactionMean phenotypic variance of F7 progeniesHeritability (%)Realized heritability (%)Expected gain with selection (%)Realized gain with selection (%)80 Abreu et al.Table IV - Mean seed yield (kg/ha) of the twenty best progenies at each location and within the locationmean.Patos de Minas Sete Lagoas MeanProgeny Yield Progeny Yield Progeny Yield6 2333 31 2031 61 209411 2333 68 1%2 111 ';17 2143 70 1%4 312 207869 2163 84 1903 60 195170 2687 93 1%1 701•2 232684 2154 125 2033 841.2 202987 2295 148 2113 932 1948118 2207 204 1982 1181 2036123 2233 208 2039 1482 1901128 2244 221 1963 1931 2068193 2335 227 1969 2082 1885221 2532 240 1955 2211.2 2248227 2367 286 2142 22i,2 2168. 239 2158 303 1946 2391 2005299 2178 323 2148 2862 1993303 2171 358 1%7 3031,2 2059309 2214 446 1910 3232 1986338 2442 458 1911 3381 2075476 2146 484 2048 4582 1774481 2194 485 1938 4811 2\<:»:1 and 2, Progeny selected at Patos de Minas and Sete Lagoas, respectively.Progeny X planting density interaction, although not significant played a con-siderable role in the estimate of genetic variance components. This was confirmed bycomparing the estimation of genetic variance among F8 progenies obtained viavariance components, in which the F8 progeny x planting density interaction is con-tained, with the same estimate obtained by the covariance between F8 progenies atthe densities of 8 and 16 plants per meter (Table III), which directly reflects geneticvariance, because no covariance exists between errors. It can be seen that the inter-action overestimated genetic variance, and consequent1y the gain obtained by selec-tion would also be overestimated if the effect of this interaction were not eliminated.Heritability estimates (h2) were obtained from genetic covariance betweenprogenies in the two environments since, as commented above, the latter directlyGenotype x Environment Interadion in Dry Beans 81reflects genetic variance without the presence of genotype x environment interaction.Narrow-sense heritability was similar to realized heritability (Table III) which was ob-tained by the gain realized with selection by simulating a selection at Patos and es-timating gain at Sete Lagoas. Similarly, the gain expected through selection waspractically identical to realized gain, further supporting the fact that estimation ofgenetic parameters through variance components, with care taken to reduce the ef-fects of genotype x environment interaction, is a powerful tool in the development ofbreeding programs.RESUMOCom o objetivo de verificar a influência da interação genótipo por ambiente na estimativa deparâmetros genéticos e fenotípicos no feijoeiro foram avaliadas 97 progênies provenientes do cruzamen-to entre as cultivares Carioca BO e Rio Tibagi,juntamente com três testemunhas ("Carioca BO", "RioTibagi"e "Carioca 1030"), quanto a produtividade de grãos, nas gerações F7 e F8' Na geração F7 a avaliação foirealizada em Patos de Minas no delineamento látice simples 10 x 10. Na geração F8o plantio foi efetuadoem Sete Lagoas, empregando-se duas densidades de semeadura, 8 e 1"6plantas por metro. Cada densidadeconstituiu um experimento para o qual adotou-se o delineamento látice simples 1.0x 10. Os resultados ob-tidos permitiram verificar a necessidade das avaliações de progênies serem efetuadas em vários ambien-tes. Tanto foi assim que a herdabilidade no sentido restrito, obtida via covariância entre as progênies nos'dois ambientes foi semelhante a herdabilidade realizada, assim como o ganho esperado com a seleção foipraticamente igual ao ganho realizado. Também a eficiência da seleção baseada na média dos ambientesfoi cerca de três vezes maior do que quando se considerou cada ambiente particular.REFERENCESA1buquerque, M.M. and Vieira, C. (1974). Manifestações da heterose em Phaseolus vulgaris L. Ciência e~Prática 21: 148-166.Beaver, J.S., Paniagua, C.V., Coyne, D.P. and Freitas, G.F. (1985). Yield stability of dry bean genotypesin the Dominican Republic. Crop. Sei. 25: 923-926.Chung, J.H. and Stevenson, E. (1973). Diallelanalyses ofthe geneticvariation in some quantitative charac-ters in dry beans. N. Z. J. Agric. Res. 16: 223-231.Fehr, W.R (1987). PrincipIes 01 cultivar development; theory and technique. MacMillan Publishing Com-pany, New York.Foolad, M.R and Bassiri, A. (1983). Estimares of combining ability, reciprocal effects and heterosis foryield and yield components in a common bean diallel cross. J. Agric. Sei. 1OQ: 103-108.Hamblin, J. and Evans, A.M. (1976). The estimation of cross yield using early generation and parentalyields in dry beans (Phaseolus vulgaris L.). Euphytica 25: 515-520.Hamblin, J. and Morton, J.R (1977). Genetic interpretations of the effects of bulk breeding on fourpopulations of beans (Phaseolus vulgaris L.). Euphytica I: 75-83.82 Abreu et al.Hamblin, J. and Zimmennann, M.J. de O. (1986). Breeding common beanfor yield in mixtures. PlantBreeding Reviews 4: 245-272.Nienhuis, J. and Singh, S.P.(1985). Effects of location and plant density on yield and architectural traitsin dry beans. Crop. Sei. 25: 579-584.Pacova, B.E.V., Candal Neto, J.F., Guidoni, AL., Santos, AF., Vargas, AA.T. and Dessaune Filho, N.D.(1987). Adaptação e estabilidade fenotípica de cultivares de feijão preto no Estado do Espírito Santo.Pesq.Agropec. Bras. 22: 491-500.Park, S.J. (1987). Cultivar by environment interactions, yield stability and grouping of test locations forfield bean cultivar trials in Ontario. Cano J. PI. Sei. 67: 653-659.Quifiones, F.A (1%9). Relationships between parents and selections in crosses of dry beans. Crop.-sci.9: 673-675.Ramalho, MA.P., Santos, J.B. and Pereira Filho, IA.P. (1988). Choise ofparents fordrybean (Phaseolusvulgaris L.) breeding. I. Interaction of mean components by generation and by location. Rev. Brasil.Gen. 11: 391-400.Rosiele, A and Hamblin, J. (1981). Theoretical aspects of selection for yield in stress and non-stress en-vironments. Crop. Sei. 21: 943-946.Santos, J.B. (1980). Estabilidade fenotípica de cultivares de feijão (Phaseotus vulgaris L.) nas condiçõesdo Sul de Minas. Masters Thesis, ESALQ, Piracicaba, SP.Santos, J.B., Vencovsky, R and Ramalho, M.AP. (1985). Controle genético da produção de grãos e deseus componentes primários em feijoeiro. Pesq. Agropcc. Bras. 20: 1203-1211.Sarafi, A (1978). A yield-component selection experiment involving arnerican and iranian cultivars of thecommon bean. Crop. Sei. 18: 5-7.Vencovsky, R (1987). Herança quantitativa. In: Melhôratnento eprodução de milho no Brasil. (Patemiani,E. and Viegas, G.P., eds.). Fundação Cargill, Campinas. pp. 135-215.Vieira, C. (1967). Ojeijoeiro-comum: cultura, doenças e melhoramento. Imprensa Universitária, Viçosa.Westennann, D.T. and Crothers, S.E. (1977). Plant population effects on lhe seed yield components ofbeans. Crop Sei. 17: 493-496.(Received Marrh 30, 1989)

Effect of genotype x environment interaction on estimations of genetic and phenotypic parameters of common beans.

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