Genetics of tassel branch number in maize and its implications for a selection program for small tassel size

Summary Tassel branch numbers of six crosses of maize (Zea mays L.) were analyzed to determine inheritance of this trait. Generation mean analyses were used to estimate genetic effects, and additive and nonadditive components of variance were calculated and evaluated for bias due to linkage. Both narrow-sense and broad-sense heritabilities were estimated. Additive genetic variance estimates were significant in five of the six crosses, whereas estimates of variance due to nonadditive components were significant in only three crosses. Additionally, estimates of additive variance components usually were larger than corresponding nonadditive components. There was no evidence for linkage bias in these estimates. Estimates of additive genetic effects were significant in four of six crosses, but significant dominance, additive × additive and additive × dominance effects also were detected. Additive, dominance, and epistatic gene action, therefore, all influenced the inheritance of tassel branch number, but additive gene action was most important. Both narrow-sense and broadsense heritability estimates were larger than those reported for other physiological traits of maize and corroborated conclusions concerning the importance of additive gene action inferred from analyses of genetic effects and variances. We concluded that selection for smalltasseled inbreds could be accomplished most easily through a mass-selection and/or pedigree-selection system. Production of a small-tasseled hybrid would require crossing of two small-tasseled inbreds. We proposed two genetic models to explain unexpected results obtained for two crosses. One model involved five interacting loci and the other employed two loci displaying only additive and additive × additive gene action.Journal Paper No. J-9231 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa 50011. Project No. 2152     

Nonadditive genetic components in resistance of the red flour beetle Tribolium castanaeum against parasite infection

Genetically coupled antagonistic coevolution between host and parasites can select for the maintenance of recombination in the host. Mechanistically, maintenance of recombination relies on epistatic interactions between resistance genes creating linkage disequilibria (LD). The role of epistasis in host resistance traits is however only partly understood. Therefore, we applied the joint scaling principle to assess epistasis and other nonadditive genetic components of two resistance traits, survival, and parasite spore load, in population crosses of the red flour beetle Tribolium castanaeum under infections with the microsporidian Nosema whitei. We found nonadditive components only in infected populations but not in control populations. The genetic architecture underlying survival under parasite infection was more complex than that of spore load. Accordingly, the observed negative correlation between survival and spore load was mainly based on a correlation between shared additive components. Breakdown of resistance was especially strong in F2 crosses between resistant lines indicating that multiple epistatic routes can lead to the same adaptation. In general, the wide range of nonoverlapping genetic components between crosses indicated that parasite resistance in T. castanaeum can be understood as a multi peaked fitness landscape with epistasis contributing substantially to phenotypic differentiation in resistance.     

Genetic basis of resistance to zonate leaf spot disease in forage sorghum

Summary Generation mean analysis was carried out for ten crosses between two resistant and two susceptible parents to find the genetic basic of resistance to zonate leaf spot disease in forage sorghum. In all crosses except one, at least one type of non-allelic interaction was present. Both additive and dominance gene effects were significant for most crosses. Duplicate type epistasis was present for the inheritance of this disease. Resistance to this disease revealed overdominance. Appropriate breeding plans were suggested to exploit the disease resistance.     

Genetic studies on resistance to Valsa canker in apple: genetic variance and breeding values estimated from intra- and inter-specific hybrid progeny populations

Malus sieboldii Rehd. exhibits high levels of resistance to Valsa canker caused by Valsa ceratosperma (Tode ex Fr.) Maire while cultivated apples (Malus domestica Borkh.) are susceptible to the disease. In this study, progenies from 23 full-sib families derived from both inter- and intra-specific hybridization among 16 Malus genotypes as parents were assessed for resistance to V. ceratosperma (Vc) for two seasons using an excised shoot assay to determine the pattern of inheritance of the resistance and to also estimate the variance components, narrow-sense heritability, and breeding values of parental genotypes. Generally, M. sieboldii × M. domestica and its reciprocal crosses had more resistant progenies to Vc than intra-specific crosses of M. domestica. Resistance to Vc expressed as the relative lesion length among progenies showed continuous variation irrespective of cross, suggesting the quantitative nature of the resistance to the three virulent isolates of Vc that were tested. Resistance to Vc using the progeny population was analyzed using a mixed linear model based on restricted maximum likelihood. The parental effect (general combining ability (GCA)) was significant while the interaction effect between parents (specific combining ability (SCA)) was relatively small and non-significant. The ratio of SCA/GCA variance was about 32%, suggesting that additive genetic variance had a major contribution to the total genetic variance for resistance to Vc. There was a positive correlation (r = 0.49, p < 0.01) between mid-parental GCA and SCA predictions among 23 full-sib families for the resistance. Narrow-sense heritability estimated by sib analysis was moderate ( [^(h)]² = 0.29 ) \left( {{{\hat{h}}^2} = 0.29} \right) . The predicted breeding values (BV) of the 16 parents indicated that M. sieboldii “Sanashi 63” and “Hayanarisanashi 1” would be useful for breeding for high levels of resistance to Vc.     

Relationship between epistasis and aggressiveness in resistance of pepper (Capsicum annuum L.) to Phytophthora nicotianae

This study evaluated the types of gene action governing the inheritance of resistance to Phytophthora nicotianae necrosis in populations derived from two crosses involving two susceptible (Beldi and Nabeul II) and one resistant (CM334) cultivars of pepper (Capsicum annuum L.). Populations, composed of Pr, Ps, F(1) , F (2) , BC (1) Pr, and BC (1) Ps generations, were inoculated with six P. nicotianae isolates. Generation means analysis indicated that an additive-dominance model was appropriate for P. nicotianae isolates Pn (Ko1) , Pn (Ko2) and Pn (Kr1) , which showed low aggressiveness in the two crosses. For the more aggressive isolates Pn (Bz1) , Pn (Bz2) and Pn (Kr2) , epistasis was an integral component of resistance in the two crosses. The presence of epistasis in the resistance of pepper to P. nicotianae was dependent on the level of aggressiveness of the isolates. Selection in pepper with less aggressive isolates was efficient, but not with more aggressive isolates; on the other hand, selection with more aggressive isolates was more stable. The minimum number of genes controlling resistance was estimated at up to 2.71. In the majority of cases, the additive variance was significant and greater than the environmental and dominance variance.     

Epistasis and the Genetic Divergence of Photoperiodism between Populations of the Pitcher-Plant Mosquito, Wyeomyia Smithii

Parallel crosses between each of two southern (ancestral) and one northern (derived) population of the pitcher-plant mosquito, Wyeomyia smithii, were made to determine the genetic components of population divergence in critical photoperiod, a phenological trait that measures adaptation to seasonality along a climatic gradient. Joint scaling tests were used to analyze means and variances of first- and second-generation hybrids in order to determine whether nonadditive genetic variance, especially epistatic variance, contributed to divergence in critical photoperiod. In both crosses, digenic epistatic effects were highly significant, indicating that genetic divergence cannot have resulted solely from differences in additively acting loci. For one cross that could be tested directly for such effects, higher order epistasis and/or linkage did not contribute to the divergence of critical photoperiod between the constituent populations.     

Evaluation of inheritance to leaf rust in wheat using area under disease progress curve

Six varieties, Kundan (K), Galvez-87 (G), Trap (T), Chris (C), Mango (M) and PBW-348 (P) along with fast ruster, Agra Local (AL), were screened for seedling reaction and adult pant response to leaf rust. Seedlings of all six varieties were susceptible while adult plants showed lower susceptability response than Agra Local. The F1s among the varieties, and also with Agra Local, showed the values lesser than the respective mid parental values for AUDPC suggesting a polygenic mode of inheritance. ANOVA for combining ability effects indicated variation due to the GCA and SCA effects, which indicated that both additive as well as non-additive type of genetic variances, govern AUDPC. The higher values for the GCA variance over the SCA variance indicated the predominance of an additive component over the dominance component for AUDPC. Significant values for GCA effects indicated that Kundan, Galvez-87 and Trap can be used as good general combiners for AUDPC. The crosses, KxAL, GxAL and TxAL showed significant sca effects for AUDPC, which indicated the predominance of non-additive gene effects in these crosses. Additive x additive and dominance x dominance components of the 5- parameter model were highly significant and contributed maximum extent compared to the additive and dominance components in the cross KxG, while dominance and dominance x dominance components contributed maximum in the remaining crosses. Under such a situation, improvement in the character may be expected through standard selection procedure, which may first exploit the additive gene effects and simultaneously care should be taken to see that the dominance effects are not dissipated, but rather they should be concentrated.     

Rice (<i>Oryza sativa</i> L.) Breeding among Hassawi Landrace and Egyptian Genotypes for Stem Borer (<i>Chilo agamemnon</i> Bles.) Resistance and Related Quantitative Traits

Rice stem borer (Chilo agamemnon Bles.) is a primary insect pest of rice and is a major limiting factor to rice production. Breeding for insect-resistant crop varieties has been an economic way of integrated pest management (IPM) as it offers a viable and ecologically acceptable approach. This study was aimed to evaluate rice genotypes for their resistance against rice stem borer. Seven parental genotypes with twenty one F1 crosses were evaluated for genotypic variation in field experiments. Analysis of variance revealed significant differences for the studied traits in almost all crosses and parents. In addition, the mean squares of parents versus their crosses were signifi- cant for stem borer resistance and other associated traits. Moreover, both general combining ability (GCA) and specific combining ability (SCA) variances were highly significant for all characters studied in the F1 generation. Based on GCA, 4 genotypes (Sakha101, Gz6903-3-4-2-1, Gz9577-4-1-1 and Hassawi) exhibited highly significant negative values for stem borer resistance (–0.53, –1.06, –0.18 and –0.49, respectively) indicating they are the best combiners for stem borer resistance. Based on SCA analysis, nine cross combinations showed highly significant negative effects for stem borer resistance. Similarly, the cross Giza178 Hassawi was the best combination with significantly highest value for early maturity. In addition, seven crosses showed highly significant negative SCA for plant height trait. On the other hand, for panicle length, number of primary branches/panicle, panicle weight and 1000-grain weight, seven, four, eight and six crosses showed highly significant positive SCA, respectively. The result further revealed that the non-additive dominance genetic variance was higher than the additive variance for all evaluated traits indicating that non-additive genetic variances have a role in their inheritance. The broad-sense heritability estimates were high for all the studied traits. The stem borer resistance was significantly correlated with panicle weight and 1000-grain weight, which also showed a highly significant correlation with grain yield/plant. Thus these traits can be effectively employed in a breeding program to confer resistance against stem borer infestation in rice. It was further supported by biplot analysis, which clustered these potentially important traits into two quadrants showing their importance in any future breeding program to control stem borer infestation. This study has contributed valuable information for evaluation of genetic diversity in the local rice germplasm and its utilization in futuristic rice genetic improvement programs.     

Inter-locus interactions: A review of experimental evidence

In quantitative genetics, experiments designed to elucidate the nature of gene action and hence the importance of epistasis, have included analysis of genetic differences among individuals in random mating populations (partitioning of genetic variation, analysis of selection responses), of differences among inbred lines or selected populations (variance components in crosses among lines, chromosome analysis using genetic markers and crossover suppression), of the effects of inbreeding, and of population structure. Evidence in population genetic studies has come from studies of linkage disequilibrium and co-adaptation in natural populations, and of multilocus fitness estimation and linkage disequilibrium and associative overdominance in experimental populations. While it is clear that epistasis does contribute to the genetic variation in some quantitative characters, and in particular reproductive fitness, much of the evidence is equivocal and unsatisfying.     

Estimation of maternal,sex-linked and additive x additive epistatic gene effects for body size of Tribolium

Summary The genetic structure of two quantitative traits, 13-day larval weight and pupal weight, in two unselected populations of Tribolium castaneum was investigated by the genetic model of Carbonell et al. (1983). The variability among two and three-way crosses was analyzed into components due to: general and specific combining abilities, maternal, sex-linkage, specific reciprocal and additive-by-additive epistasis. Also, indirect evidence of higher order epistasis was studied. It is concluded that the heterotic trait larval weight is highly affected by sex-linked genes and by non-additive gene action with additive-by-additive as well as higher order epistasis playing major roles. Pupal weight, on the other hand, is determined mostly by additive gene action although epistasis is also a significant source for genetic variability. Both traits are significantly influenced by maternal effects.Journal Paper No. 9033 from Purdue University Agricultural Experiment Station. Based in part on PhD Theses submitted by the first two authors at Purdue University     

Population differentiation in the beetleTribolium castaneum. I. Genetic architecture

We used joint-scaling analyses in conjunction with rearing temperature variation to investigate the contributions of additive, non-additive, and environmental effects to genetic divergence and incipient speciation among 12 populations of the red flour beetle, Tribolium castaneum, with small levels of pairwise nuclear genetic divergence (0.033 < Nei's D < 0.125). For 15 population pairs we created a full spectrum of line crosses (two parental, two reciprocal F1's, four F2's, and eight backcrosses), reared them at multiple temperatures, and analyzed the numbers and developmental defects of offspring. We assayed a total of 219,388 offspring from 5147 families. Failed crosses occurred predominately in F2's, giving evidence of F2 breakdown within this species. In all cases where a significant model could be fit to the data on offspring number, we observed at least one type of digenic epistasis. We also found maternal and cytoplasmic effects to be common components of divergence among T. castaneum populations. In some cases, the most complex model tested (additive, dominance, epistatic, maternal, and cytoplasmic effects) did not provide a significant fit to the data, suggesting that linkage or higher order epistasis is involved in differentiation between some populations. For the limb deformity data, we observed significant genotype-by-environment interaction in most crosses and pure parent crosses tended to have fewer deformities than hybrid crosses. Complexity of genetic architecture was not correlated with either geographic distance or genetic distance. Our results support the view that genetic incompatibilities responsible for postzygotic isolation, an important component of speciation, may be a natural but serendipitous consequence of nonadditive genetic effects and structured populations.     

WILL POPULATION BOTTLENECKS AND MULTILOCUS EPISTASIS INCREASE ADDITIVE GENETIC VARIANCE?

We apply new analytical methods to understand the consequences of population bottlenecks for expected additive genetic variance. We analyze essentially all models for multilocus epistasis that have been numerically simulated to demonstrate increased additive variance. We conclude that for biologically plausible models, large increases in expected additive variance--attributable to epistasis rather than dominance--are unlikely. Naciri-Graven and Goudet (2003) found that as the number of epistatically interacting loci increases, additive variance tends to be inflated more after a bottleneck. We argue that this result reflects biologically unrealistic aspects of their models. Specifically, as the number of loci increases, higher-order epistatic interactions become increasingly important in these models, with an increasing fraction of the genetic variance becoming nonadditive, contrary to empirical observations. As shown by Barton and Turelli (2004), without dominance, conversion of nonadditive to additive variance depends only on the variance components and not on the number of loci per se. Numerical results indicating that more inbreeding is needed to produce maximal release of additive variance with more loci follow directly from our analytical results, which show that high levels of inbreeding (F > 0.5) are needed for significant conversion of higher-order components. We discuss alternative approaches to modeling multilocus epistasis and understanding its consequences.     

The contrasting genetic architecture of wing size, viability, and development time in a rainforest species and its more widely distributed relative

Divergence among populations can occur via additive genetic effects and/or because of epistatic interactions among genes. Here we use line-cross analysis to compare the importance of epistasis in divergence among two sympatric Drosophila species from eastern Australia, one (D. serrata) distributed continuously and the other (D. birchii) confined to rainforest habitats that are often disjunct. For D. serrata, crosses indicated that development time and wing size differences were due to additive genetic effects, while for viability there were digenic epistatic effects. Crosses comparing geographically close populations as well as those involving the most geographically distant populations (including the southern species border) revealed epistatic interactions, whereas crosses at an intermediate distance showed no epistasis. In D. birchii, there was no evidence of epistasis for viability, although for development time and wing size there was epistasis in the cross between the most geographically diverged populations. Strong epistasis has not developed among the D. birchii populations, and this habitat specialist does not show stronger epistasis than D. serrata. Given that epistasis has been detected in crosses with other species from eastern Australia, including the recently introduced D. melanogaster, the results point to epistasis not being directly linked to divergence times among populations.     

Combining ability analysis over environments in diallel crosses of linseed (Linum usitatissimum L.)

Summary This paper reports on combining ability studies for yield and its component traits in diallel crosses involving ten ecogeographically and genetically diverse linseed (Linum usitatissimum L.) cultivars in the F₂ generation over three locations. The general combining ability (GCA) and specific combining ability (SCA) mean squares were significant at all three locations for all traits. Combined analysis over locations showed the same trend of significance. The ratio of GCA to SCA mean squares was significant for all the traits in individual location analysis as well as in combined analysis. This indicated the predominant role of additive gene effects in the inheritance of these characters. The GCA mean squares were several times larger than SCA mean squares for all the traits, indicating the presence of considerable magnitude of additive genetic variance and the additive x additive components of the epistatic variance. Consequently, effective selection should be possible within these F₂ populations for all characters. Significant genotype-location and GCA-location interactions indicated that more than one test location is required to obtain reliable information. The inexpensive and reliable procedure used for making the choice of parents was the determination of breeding values of the parents on the relative performance of their F₂ progeny bulks.Part of the thesis submitted by the senior author in partial fulfillment of the requirements for the Ph. D. degree of the Marathwada Agricultural University, Parbhani, 431 402, India     

Combined genetic analysis of partial blast resistance in an upland rice population and recurrent selection for line and hybrid values

The CNA-IRAT 5 upland rice population has been improved for 4 years by recurrent selection for blast resistance in Brazil. In order to predict the efficiency of recurrent selection in different test systems and to compare the relative advantage of hybrids versus pure line breeding, a combined genetic analysis of partial blast resistance in the CNA-IRAT 5 population was undertaken. A three-level hierarchical design in inbreeding and a factorial design were derived from the base population. Partial blast resistance of lines and hybrids was evaluated in the greenhouse and in the field by inoculation with one virulent blast isolate. The means and genetic variances of the hybrids and lines were estimated. Genetic advance by recurrent selection was predicted from estimates of variance components. The inheritance of partial blast resistance was mainly additive but non-additive effects were detected at both levels of means and variances. Mean heterosis ranged from 4%–8% for lesion size and lesion density to 10–12% for leaf and panicle resistance. High dominance or homozygous dominance variances relative to additive variance and negative covariance between additive and homozygous dominance effects were estimated. A low frequency of favourable alleles for partial resistance would explain the observed organisation of genetic variability in the base population. Recurrent selection will efficiently improve partial blast resistance of the CNA-IRAT 5 population. Genetic advance for line or hybrid values was expected to be higher testing doubled haploid lines than S1 lines, or than general combining ability. Two components of partial resistance assessed in the greenhouse, lesion size and lesion density, could be used as indirect selection criteria to improve field resistance. On the whole, hybrid breeding for partial blast resistance appeared to be slightly more advantageous than pure line breeding.     

Inheritance of photoperiod-induced flowering in three photoperiodic lines of Aeschynomene americana L.

Summary Induction of flowering by photoperiod was studied in the parental, F₁, F₂, and reciprocal backcross generations of crosses between three photoperiod-responsive Aeschynomene americana L. lines. Generation means appeared additive. Analysis with Mather and Jinks' scaling tests showed little or no epistasis and indicated that an additive-dominance model was adequate. Partitioning components of variation revealed that nearly all variation was additive genetic with dominance and environmental variation negligible. An additive genetic model with two loci, each with two alleles and all alleles having equal net effect, was tested using Power's partitioning method. Results demonstrated that the model fit the data and that there is a major additive genetic system controlling flowering in these crosses, with minor genetic and environmental influences present. Selection for flowering at a desired day length should be feasible.Accepted by A.R. HallauerFlorida Agricultural Experiment Station, Journal Series No. 9251     

THE GENETIC ARCHITECTURE OF WING SIZE DIVERGENCE AT VARYING SPATIAL SCALES ALONG A BODY SIZE CLINE IN DROSOPHILA MELANOGASTER

Latitudinal clines in quantitative traits are common, but surprisingly little is known about the genetic bases of these divergences and how they vary within and between clines. Here, we use line‐cross analysis to investigate the genetic architecture of wing size divergences at varying spatial scales along a body size cline in Drosophila melanogaster. Our results revealed that divergences in wing size along the cline were due to strong additive effects. Significant nonadditive genetic effects, including epistasis and maternal effects, were also detected, but they were relatively minor in comparison to the additive effects and none were common to all crosses. There was no evidence of increased epistasis in crosses between more geographically distant populations and, unlike in previous studies, we found no significant dominance effects on wing size in any cross. Our results suggest there is little variation in the genetic control of wing size along the length of the Australian cline. They also highlight marked inconsistencies in the magnitude of dominance effects across studies, which may reflect different opportunities for mutation accumulation while lines are in laboratory culture.     

Additive inheritance of resistance to pod rot caused by Phytophthora palmivora in cocoa

Summary Quantitative inheritance of resistance to Phytophthora pod rot (Ppr) was studied in cocoa hybrid progeny from 12 Trinitario x Amazonian crosses and their reciprocal crosses. The crossing scheme was similar to a factorial design. Disease was assessed by the number and percentage of infected pods on each tree. Highly significant differences due to general combining abilities (GCA) were obtained for all characters, except for the GCA of Trinitario on total pod production. Differences for specific combining ability (SCA) were not significant for all characters. There were no significant differences between reciprocal crosses. The Trinitario clone K82 provided the only source for the hybrid progenies of strong Ppr resistance to the hybrid progenies, while K20 provided moderate resistance. Other parental clones — KA2-101, KA5-201, KEE 2, KEE 5, and KEE 52 — produced progenies which were susceptible to Ppr. It is evident that resistance to Ppr in cocoa is inherited additively. Maternal and cytoplasmic effects were assumed to have no influence on inheritance of resistance. It is also concluded that resistance to Ppr of the kind shown by K82 is likely to be horizontal resistance. Breeding for high-yielding cultivars combined with Ppr resistance is the most effective way of controlling Ppr of cocoa on the crops of growers with small holdings in Papua New Guinea.     

Amplified fragment length polymorphism mapping of quantitative trait loci for malaria parasite susceptibility in the yellow fever mosquito Aedes aegypti

The yellow fever mosquito Aedes aegypti has been the subject of extensive genetic research due to its medical importance and the ease with which it can be manipulated in the laboratory. A molecular genetic linkage map was constructed using 148 amplified fragment length polymorphism (AFLP) and six single-strand conformation polymorphism (SSCP) markers. Eighteen AFLP primer combinations were used to genotype two reciprocal F2 segregating populations. Each primer combination generated an average of 8.2 AFLP markers eligible for linkage mapping. The length of the integrated map was 180.9 cM, giving an average marker resolution of 1.2 cM. Composite interval mapping revealed a total of six QTL significantly affecting Plasmodium susceptibility in the two reciprocal crosses of Ae. aegypti. Two common QTL on linkage group 2 were identified in both crosses that had similar effects on the phenotype, and four QTL were unique to each cross. In one cross, the four main QTL accounted for 64% of the total phenotypic variance, and digenic epistasis explained 11.8% of the variance. In the second cross, the four main QTL explained 66% of the variance, and digenic epistasis accounted for 16% of the variance. The actions of these QTL were either dominance or underdominance. Our results indicated that at least three new QTL were mapped on chromosomes 1 and 3. The polygenic nature of susceptibility to P. gallinaceum and epistasis are important factors for significant variation within or among mosquito strains. The new map provides additional information useful for further genetic investigation, such as identification of new genes and positional cloning.     

Genomic prediction of hybrid crops allows disentangling dominance and epistasis

We revisited, in a genomic context, the theory of hybrid genetic evaluation models of hybrid crosses of pure lines, as the current practice is largely based on infinitesimal model assumptions. Expressions for covariances between hybrids due to additive substitution effects and dominance and epistatic deviations were analytically derived. Using dense markers in a GBLUP analysis, it is possible to split specific combining ability into dominance and across-groups epistatic deviations, and to split general combining ability (GCA) into within-line additive effects and within-line additive by additive (and higher order) epistatic deviations. We analyzed a publicly available maize data set of Dent × Flint hybrids using our new model (called GCA-model) up to additive by additive epistasis. To model higher order interactions within GCAs, we also fitted “residual genetic” line effects. Our new GCA-model was compared with another genomic model which assumes a uniquely defined effect of genes across origins. Most variation in hybrids is accounted by GCA. Variances due to dominance and epistasis have similar magnitudes. Models based on defining effects either differently or identically across heterotic groups resulted in similar predictive abilities for hybrids. The currently used model inflates the estimated additive genetic variance. This is not important for hybrid predictions but has consequences for the breeding scheme—e.g. overestimation of the genetic gain within heterotic group. Therefore, we recommend using GCA-model, which is appropriate for genomic prediction and variance component estimation in hybrid crops using genomic data, and whose results can be practically interpreted and used for breeding purposes.