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A Review of the Evidence - Search CIMMYT repository

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est in most studies, it may be dangerous to conclude that traits such as vigor and yield areexpressed only in additive fashion. Cancellation <strong>of</strong> inter-locus interactions (epistasis) mayprevent detection <strong>of</strong> non-additive variation.Unlike most previous researchers who had focused on crosses made among inbred lines,Pollak et al. (1957) studied heterosis and inbreeding depression in crosses made among threeOPVs. In <strong>the</strong>ir study, <strong>the</strong> F1, F2, and backcross populations <strong>of</strong> <strong>the</strong> OPVs were used toestablish different levels <strong>of</strong> heterozygosity. After determining that <strong>the</strong> mean yield <strong>of</strong> <strong>the</strong> F2plants lay in between <strong>the</strong> mean yield <strong>of</strong> <strong>the</strong> parents and <strong>the</strong> mean yield <strong>of</strong> <strong>the</strong> F1 plants,Pollak et al. concluded that <strong>the</strong>ir results were consistent with an additive genetic model (withdominance) and that epistatic effects were minimal.Shehata and Dhawan (1975) evaluated inbreeding depression among three sets <strong>of</strong> diallelcrosses; each set included 10 parents that were ei<strong>the</strong>r OPVs or syn<strong>the</strong>tics. In comparing <strong>the</strong>yields <strong>of</strong> F1 and F2 plants, <strong>the</strong> average inbreeding depression for each <strong>of</strong> <strong>the</strong> three diallelswas estimated to be 10.2%, 11.4%, and 11.2%. The deviation in yield performance <strong>of</strong> <strong>the</strong> F2plants from <strong>the</strong> mean performance <strong>of</strong> <strong>the</strong> parental varieties and <strong>the</strong> F1 variety crosses wassmall in all instances, suggesting largely additive effects among loci, with limited netepistatic␣ef fects.Most studies in <strong>the</strong> plant breeding literature confirm that <strong>the</strong> arithmetic model best predicts<strong>the</strong> decline in yield performance observed between F1 and F2 plants. In a comprehensivereview, Hallauer and Miranda (1989) conclude that <strong>the</strong> yield decreases observed when F1plants are self-pollinated (inbred) to produce F2 plants generally are smaller when <strong>the</strong> F1plants are OPVs, ra<strong>the</strong>r than inbred lines. While epistatic effects are generally minimal, <strong>the</strong>ytend to be greater in line-based crosses compared to variety-based crosses. Hallauer andMiranda remark, “This difference seems reasonable because <strong>of</strong> <strong>the</strong> nature <strong>of</strong> <strong>the</strong> materialincluded in <strong>the</strong> crosses. For varieties this would involve an array <strong>of</strong> genotypes included in <strong>the</strong>crosses, whereas crosses <strong>of</strong> inbred lines would involve <strong>the</strong> expression <strong>of</strong> two specificgenotypes.” Extrapolating from this observation, one would expect that <strong>the</strong> inbreedingdepression observed in three-way crosses, double crosses, and non-conventional hybridsformed using one inbred line to fall somewhere in between that observed in single crossesand variety crosses.Ma<strong>the</strong>r and Jinks (1971) propose equations derived from Wright’s original equation to predict<strong>the</strong> performance <strong>of</strong> F2, F3, F4, and subsequent generations <strong>of</strong> plants in self-pollinated crops.For quantitative traits, <strong>the</strong> mean <strong>of</strong> F2 generation plants (obtained by selfing or sib-mating F1plants) can be calculated as follows:where:F2 = (1/4) (P1 + P2 + 2F1) (2)F2 = <strong>the</strong> mean <strong>of</strong> F2 generation plants,F1 = <strong>the</strong> cross-mean F1 generation plants,P1 = <strong>the</strong> mean <strong>of</strong> <strong>the</strong> first parent line (assumed to be completely homozygous), andP2 = <strong>the</strong> mean <strong>of</strong> <strong>the</strong> second parent line (assumed to be completely homozygous).35

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