Genetic analysis of agronomic characters in chickpea

Summary Twenty-eight diallel trials over 8 years and two locations were analysed to estimate genetic variances for agronomic characters of chickpea (Cicer arietinum L.). The data were analysed according to Method 4 and Model I of Griffing (1956). Days to flowering, plant height, and seed size were found to be predominantly under additive inheritance and were highly predictable. Both additive and non-additive genetic components were important for seed yield, number of branches, pods per plant, and seeds per pod. Although both general combining ability (gca) and specific combining ability (sca) varied significantly with generation, components of gca mean squares were invariably much larger than gca x generation interaction components, indicating that either the F₁ or the F₂ generation can be used to estimate the gca components effectively. Combined diallel analysis of F₂s over locations revealed the importance of combining ability x location interactions and emphasized the need for testing over more than one location for the precise estimation of combining ability. The implications of these findings and those reported earlier in the literature on the breeding strategies/methods for the genetic improvement of agronomic characters in chickpea are discussed.ICRISAT journal article no. 1199     

Generation mean analyses for flowering and maturity in Indian mustard (Brassica junces (L.) Czern & Coss)

Summary Components of generation means were partitioned for days to flower initiation and maturity in three crosses of Indian mustard (Brassica juncea (L.) Czern and Coss) cultivars. A linked digenics model was adequate for flowering in cross II and maturity in Cross I. All three types of digenic interactions among the linked pairs of genes, additive X additive (i), additive X dominance (j) and dominance X dominance (1), contributed significantly in the inheritance of flowering in cross II and maturity in cross I. A complete association among the genes of greater effects in higher mean parent was detected for flowering in cross II and maturity in Cross I. Duplicate epistasis was evident for flowering in Crosses I and II and maturity in Crosses I and III.Inadequacy of all the fitted models for days to flowering in Cross III and maturity in Cross II indicated the presence of higher order interactions.Part of PhD (plant breeding) Thesis, submitted by senior author to GBPUAT, Pantnagar (Nainital) U.P., India (unpublished). Research paper No. 4262     

Genetic analysis and mapping of genes controlling freezing tolerance in oilseedBrassica

Freezing tolerance is the ability of plants to survive subfreezing temperatures and is a major component of winter survival. In order to study the genetic regulation of freezing tolerance, an F2 population ofBrassica rapa and a doubled haploid population ofBrassica napus were assayedin vitro for relative freezing tolerance of acclimated and nonacclimated plants. Linkage maps developed previously were used to identify putative quantitative trait loci (QTL). Genomic regions with significant effects on freezing tolerance were not found for theB. napus population, but forB. rapa four regions were associated with acclimated freezing tolerance (FTA) and acclimation ability (FTB), and two unliked regions were associated with nonacclimated freezing tolerance (FTN). Acclimation ability was regulated by genes with very small additive effects and both positive and negative dominance effects. The allele from the winter parent at the FTN QTL had positive additive effects, but negative dominance effects. RFLP loci detected by a cold-induced and a stress-related cDNA fromArabidopsis thaliana mapped near two QTL for FTA/FTB. Further tests are needed to determine if alleles at these loci are responsible for the QTL effects we detected.     

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     

Genetic analysis of freezing tolerance in blueberry (Vaccinium section Cyanococcus)

An understanding of the genetic control of freezing tolerance (FT) in woody perennials is important for the effective selection and development of plants with a broader climatic adaptation. This study was undertaken to examine the inheritance and gene action of FT in segregating populations of a woody perennial blueberry (Vaccinium, section Cyanococcus). Two backcross populations were derived from interspecific hybrids of the diploid species Vaccinium darrowi and Vaccinium caesariense, which are widely divergent in their FT. The bud FTs of uniformly cold acclimated plants of parental, F₁, and two backcross populations were evaluated with a laboratory controlled freeze-thaw regime, followed by a visual assessment of injury. FT (LT₅₀) was defined as the temperature causing 50% of the flower buds to be injured. Data indicate that the two parents were homozygous for genes for low or high FT. Freezing-tolerance values of the parental and F₁ populations indicate that freeze-sensitivity is a partially dominant trait. Results from reciprocal crosses revealed that there was no significant maternal influence on freezing tolerance. Parental phenotypes were fully recovered in 40–42 plants of each testcross population, suggesting that FT is determined by relatively few genes. The degree of dominance and an analysis of generation means revealed that FT in blueberry is controlled largely by additive gene effects and, to a lesser degree, by dominance gene effects. Testing of various genetic models indicated that FT inheritance can be adequately explained by a simple additive-dominance model; however, two epistatic models involving additive-additive and dominance-dominance interactions also fit the data. Received: 25 May 1999 / Accepted: 16 June 1999     

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     

Genetics of leaf blight resistance in wheat

Summary Studies on the genetics of leaf blight caused byAlternaria triticina using generation mean analysis revealed that additive components played a major role, but that dominance components also contributed significantly in controlling the variability for leaf blight resistance in wheat crosses. Furthermore, the additive x additive type of epistasis was predominant in the first three crosses, whereas in the fourth cross additive x dominance (j) and dominance x dominance (1) components of epistasis were most significant. Because of this it may be desirable to follow a simple recurrent selection scheme for higher tolerance, to isolate resistant plants from the segregating populations derived from crosses of parents of diverse origin following the pedigree method of breeding. CPAN-1887 was very tolerant to leaf blight in the present study and should be utilized in hybridization programs to develop leaf-blight-resistant varieties.     

Genetic analysis of tolerance for phosphorous deficiency in rice (Oryza sativa L.)

The inheritance of phosphorous (P) — deficiency tolerance in rice was investigated by a sevenparent diallel. The parent materials involved were four P-efficient (IR20, IR54, IR28, and Mahsuri), one moderately P-efficient (TN1), and two P-inefficient (IR31406333-1 and IR34686-179-1-2-1), genotypes. Relative tilering ability (RTA) under P-deficient and P-supplemented soil conditions was the parameter used in determining the tolerance level of the different genotypes. Diallel graph analysis revealed that tolerant parents have an excess of recessive genes, while moderate and susceptible parents possess more dominant genes. Genetic-component analysis suggested that both additive and dominance gene effects are involved in the inheritance of P-deficiency tolerance in rice. The trait exhibited over doiminance as confirmed by the graphical analysis. Narrow-sense heritability of the trait was moderate (0.50) and environmental effects were low. Both the general combining ability (GCA) and the specific combining ability (SCA) were significant, but GCA was more prevalent than SCA. Tolerant parents exhibited a high GCA whereas susceptibles have a very poor GCA, suggesting that tolerant parents were mostly enriched in additive genes and susceptible parents in non-additive genes. Crosses involving two high general combiners showed low SCA effects whereas crosses between poor general combiners manifested highly-significant SCA values.     

Epistasis in maize (Zea mays L.)

Summary Three flint and three dent maize (Zea mays L.) inbred lines, their possible F₁ crosses, F₂ and backcross progenies, and all possible three-way crosses were evaluated in a three-year experiment for yield, ear moisture, and plant height. The purpose was to estimate genetic parameters in European breeding materials from (i) generation means analysis, (ii) diallel analysis of generation means, and (iii) analysis of F₁ and three-way cross hybrids. Method (i) was based on the F-metric model and methods (ii) and (iii) on the Eberhart-Gardner (1966) genetic model; both models extended for heterotic maternal effects.Differences among generation means for yield and plant height were mainly attributable to dominance effects. Epistatic effects were significantly different from zero in a few crosses and considerably reduced heterosis in both traits. Additive x additive and domiance x dominance effects for yield were consistently positive and negative, respectively. Significant maternal effects were established to the advantage of generations with a heterozygous seed parent. In the diallel analysis, mean squares for dominance effects were greater than for additive effects for yield and plant height but smaller for ear moisture. Though significant for yield and plant height, epistatic variation was small compared to additive and dominance variation. Estimates of additive x additive epistasis for yield were significantly negative in 11 of 15 crosses, suggesting that advantageous gene combinations in the lines had been disrupted by recombination in the segregating generations. The analysis of hybrids supported the above findings regarding the analysis of variance. However, the estimates of additive x additive epistasis for yield were considerably smaller and only minimally correlated with those from the diallel analysis. Use of noninbred materials as opposed to materials with different levels of inbreeding is considered the main reason for the discrepancies in the results.     

Gene actions at loci underlying several quantitative traits in two elite rice hybrids

To understand the gene activities controlling nine important agronomic quantitative traits in rice, we applied a North Carolina design 3 (NC III design) analysis to recombinant inbred lines (RILs) in highly heterotic inter- (IJ) and intra-subspecific (II) hybrids by performing the following tasks: (1) investigating the relative contribution of additive, dominant, and epistatic effects for performance traits by generation means analysis and variance component estimates; (2) detecting the number, genomic positions, and genetic effects of QTL for phenotypic traits; and (3) characterizing their mode of gene action. Under an F∞-metric, generation means analysis and variance components estimates revealed that epistatic effects prevailed for the majority of traits in the two hybrids. QTL analysis identified 48 and 66 main-effect QTL (M-QTL) for nine traits in IJ and II hybrids, respectively. In IJ hybrids, 20 QTL (41.7%) showed an additive effect of gene actions, 20 (41.7%) showed partial-to-complete dominance, and 8 (16.7%) showed overdominance. In II hybrids, 34 QTL (51.5%) exhibited additive effects, 14 (21.2%) partial-to-complete dominance, and 18 (27.3%) overdominance. There were 153 digenic interactions (E-QTL) in the IJ hybrid and 252 in the II hybrid. These results suggest that additive effects, dominance, overdominance, and particularly epistasis attribute to the genetic basis of the expression of traits in the two hybrids. Additionally, we determined that the genetic causes of phenotypic traits and their heterosis are different. In the plants we studied, the phenotypic traits investigated and their heterosis were conditioned by different M-QTL and E-QTL, respectively, and were mainly due to non-allelic interactions (epistasis).     

Analysis of diallel cross in Phaseolus aureus roxb

Summary In a diallel cross of Phaseolus aureus involving five varieties, combining ability and gene action for grain yield, grains per pod and pods per plant were estimated. The study indicated that both general combining ability and specific combining ability effects were significant and important for all three traits. Partial dominance for grain yield and partial to over-dominance for grains per pod and pods per plant were observed. Dominant genes seem to govern the inheritance of all three characters. Combining ability, and graphical and component variance analyses indicated that the grain yield and two of its components are influenced by both additive and non-additive gene action.     

Combining ability analysis over F1-F5 generations in diallel crosses of bread wheat

Summary Combining ability studies for grain yield and its primary component traits in diallel crosses involving seven diverse wheat cultivars of bread wheat (Triticum aestivum L.) over generations F₁-F₅ are reported. The general and specific combining ability variances were significant in all generations for all the traits except specific combining ability variance for number of spikes per plant in the F₅. The ratio of general to specific combining ability variances was significant for all the traits except grain yield in all the generations. This indicated an equal role of additive and non-additive gene effects in the inheritance of grain yield, and the predominance of the former for its component traits. The presence of significant specific combining ability variances in even the advanced generations may be the result of an additive x additive type of epistasis or evolutionary divergence among progenies in the same parental array. The relative breeding values of the parental varieties, as indicated by their general combining ability effects, did not vary much over the generations. The cheap and reliable procedure observed for making the choice of parents, selecting hybrids and predicting advanced generation (F₅) bulk hybrid performance was the determination of breeding values of the parents on the relative performance of their F₂ progeny bulks.     

Inheritance of partial resistance to powdery mildew in spring wheat

Summary Four spring wheat (Triticum aestivum L.) cultivars exhibiting partial resistance to powdery mildew induced by Erysiphe graminis f.sp. tritici were crossed to a common susceptible cultivar to study the inheritance of resistance. The genetic parameters contributing to resistance were estimated by generation means analyses. Additive gene action was the most important genetic component of variation among generation means in all four crosses. Additive by additive effects were significant in one cross and both additive by additive and additive by dominance effects were significant in another. Dominance effects were not significant. The F2/F3 correlations in three crosses ranged from 0.27 to 0.43. Three additional crosses among resistant cultivars were employed to study the effectiveness of selection in improving resistance. By selecting the most resistant plants from the F2 and evaluating the progenies in the F4, increases in resistance ranging from 21% to 31% were obtained. In all crosses, there was transgressive segregation in both directions indicating that the genes conferring resistance to these cultivars differ and exhibit additive effects.     

Estimation of genetic parameters in barley (Hordeum vulgare L.)

Summary Four exotic and four indigenous strains of barley were used for making diallel crosses. The sets of parents and crosses making full, half and quarter diallel were analysed in a randomized block design for plant height, number of effective tillers, ear length, grain yield per plant, 100 grain weight and number of grains per ear.The three alternatives of diallel were similar with respect to the estimates of degree of dominance, general combining ability and specific combining ability, indicating that all these three methods of diallel were equally efficient. However, as the number of entries are minimum in quarter diallel, it would be economical in terms of cost, time and labour to estimate genetic parameters by this method. Average degree of dominance was found in the range of overdominance. The ranking of parents on the basis of their array mean was similar to the ranking based on gca effects. Similarly, the ranking of crosses on the basis of per se performance was similar to the ranking based on sca effects. This suggests that the selection of best general combiner or best cross combinations may be easier and more effective through array mean for per se performance rather than through high gca and sea effects, respectively. From among 56 crosses, IB-226 X X C-164 was the one which showed superiority for maximum number of characters followed by AB-12/59 X PTS-57. High sea effect for plant height, ear length, grain yield, 100 grain weight and number of grains per ear was the result of cross between parents having high X low general combining ability, indicating additive X dominance type of gene interaction. For number of effective tillers, high sca was produced by low x low general combiners, indicating dominance x dominance gene interaction.Part of a Ph. D. thesis submitted by senior author to Haryana Agricultural University, Hissar.     

Inheritance of deeper root length and grain yield in half-diallel durum wheat (Triticum durum) crosses

Inheritance, heterosis and combining ability of deeper root length (DRL) and grain yield (GY) were investigated in durum wheat populations obtained from half‐diallel crossings among five parental lines differing in their DRL and GY. The study was conducted with the final objective of identifying parent lines to be used in a breeding programme to develop drought‐tolerant wheat varieties. General combining ability and specific combining ability effects were significant for both traits; however, additive gene effects were predominant over non‐additive effects. Partial dominance was ambidirectional for DRL and unidirectional for GY. Lines INRAT69 and Omrabia conferred DRL whereas Omrabia and Khiar transmitted high GY to their respective progenies. In the studied material, both characters were controlled mainly by dominant alleles, but they could also be attributed to recessive alleles although less frequently. Both broad‐sense and strict‐sense heritabilities were high for DRL, confirming the importance of additive gene effects, whereas strict‐sense heritability for GY was average, indicating the importance of interaction effects as compared with the additive effects; this could mean reduced selection efficiency for the latter trait. Thus, the expected genetic progress per cycle of selection will be lower for GY compared with DRL. Omrabia should be included in the breeding programme as a parent so that while maintaining high GY, resulting progeny should be better able to resist drought through DRL.     

Local Adaptation and Genetics of Acid-Stress Tolerance in the Moor Frog, Rana arvalis

As potential to adapt to environmental stress can be essential for population persistence, knowledge on the genetic architecture of local adaptation is important for conservation genetics. We investigated the relative importance of additive genetic, dominance and maternal effects contributions to acid stress tolerance in two moor frog (Rana arvalis) populations originating from low and neutral pH habitats. Experiments with crosses obtained from artificial matings revealed that embryos from the acid origin population were more tolerant to low pH than embryos from the neutral origin population in embryonic survival rates, but not in terms of developmental stability, developmental and growth rates. Strong maternal effect and small additive genetic contributions to variation were detected in all traits in both populations. In general, dominance contributions to variance in different traits were of similar magnitude to the additive genetic effects, but dominance effects outweighed the additive genetic and maternal effects contributions to early growth in both populations. Furthermore, the expression of additive genetic variance was independent of pH treatment, suggesting little additive genetic variation in acid stress tolerance. The results suggest that although local genetic adaptation to acid stress has taken place, the current variation in acid stress tolerance in acidified populations may owe largely to non-genetic effects. However, low but significant heritabilities (h ² 0.07–0.22) in all traits – including viability itself – under a wide range of pH conditions suggests that environmental stress created by low pH is unlikely to lower moor frog populations' ability to respond to selection in the traits studied. Nevertheless, acid conditions could lower populations' ability to respond to selection in the long run through reduction in effective population size.     

Identification of X-linked quantitative trait loci affecting cold tolerance in Drosophila melanogaster and fine mapping by selective sweep analysis

Drosophila melanogaster is a cosmopolitan species that colonizes a great variety of environments. One trait that shows abundant evidence for naturally segregating genetic variance in different populations of D. melanogaster is cold tolerance. Previous work has found quantitative trait loci (QTL) exclusively on the second and the third chromosomes. To gain insight into the genetic architecture of cold tolerance on the X chromosome and to compare the results with our analyses of selective sweeps, a mapping population was derived from a cross between substitution lines that solely differed in the origin of their X chromosome: one originates from a European inbred line and the other one from an African inbred line. We found a total of six QTL for cold tolerance factors on the X chromosome of D. melanogaster. Although the composite interval mapping revealed slightly different QTL profiles between sexes, a coherent model suggests that most QTL overlapped between sexes, and each explained around 5–14% of the genetic variance (which may be slightly overestimated). The allelic effects were largely additive, but we also detected two significant interactions. Taken together, this provides evidence for multiple QTL that are spread along the entire X chromosome and whose effects range from low to intermediate. One detected transgressive QTL influences cold tolerance in different ways for the two sexes. While females benefit from the European allele increasing their cold tolerance, males tend to do better with the African allele. Finally, using selective sweep mapping, the candidate gene CG16700 for cold tolerance colocalizing with a QTL was identified.     

Inheritance of resistance to wheat yellow rust at adult plant stage

In order to investigate on inheritance and gene action for resistance to yellow rust, the resistant line C.B227 was crossed with the susceptible variety Avocet. Parents (P1 and P2) and the resulting F1, F2 and F3 generations were planted in a randomised complete block design with two replications in the field. The plants were inoculated with 70E0A⁺ pathotype of yellow rust in the research station of Gharakhil, Iran, and evaluated for resistance at adult plant stage. Disease severity and infection type of flag leaf were recorded for each single plant and final coefficient of infection was calculated. The results of weighted ANOVA indicated that the difference among the generations was significant (p?<?0.01) for the trait final infection type. Generation mean analysis showed that dominant effect was more important than additive one. The degree of dominance indicated the presence of complete dominance. Additive, dominance and epistasic additive?×?additive [i] effects were important in genetic control of resistance. The results of generation variance analysis were consistent with generation mean analysis.     

Genetic analyses for certain plant and ear characters in pearl millet top-crosses

Summary Combining ability and the genetics of tiller number, days taken to flower and ear thickness were studied in top-cross progenies of pearl millet. General combining ability seemed to be more important for all the characters. The prevalence of epistatic variation, presumably of the type additive x additive, additive x dominance and dominance x dominance gene effects, with a non-significant contribution of additive and dominance components of genetic variance, was observed for tiller number. For days taken to flower, the importance of additive genetic variance was greater than that of the dominance component with directional dominance towards the recessive allele. However, for ear thickness, the existence of additive genetic variability together with the additive x additive type of genic interaction was suggested.An appreciable effect of epistasis on and ₁ components was observed for tiller number, whereas this effect was not so marked for other characters.The author is grateful to Dr. D. S. Athwal, formerly Professor and Head (now Assistant Director, The International Rice Research Institute, Los Banõs, Laguna, Philippines), Department of Plant Breeding, Punjab Agricultural University, Ludhiana, for providing the facilities.     

Generating data in models including direct and maternal dominance effects

Algorithms are presented to simulate multiple generations of animal data by a model including direct additive genetic, maternal additive genetic, direct dominance, maternal dominance and permanent environmental effects. Dominance effects were computed as parental subclasses. Testing involved five single trait models that included direct contemporary group and direct additive effects, and different combinations of maternal, permanent environmental, and dominance effects. Simulated populations included 5 generations of animals and 20 contemporary groups per generation. The base population contained 200 sires and 600 dams. Variance components were estimated by Average-Information Restricted Maximum Likelihood (AIREML). No significant bias was observed. The simulation algorithms can be used in research involving dominance models, such as evaluation of mating systems exploiting special combining abilities of prospective parents.