Combining ability in the F1 and F2 generations of diallel cross in hexaploid wheat (Triticum aestivum L. em. Thell)

The F(1) and F(2) progenies of a ten-parent diallel cross (excluding reciprocals) of hexaploid wheat (Triticum aestivum L. em. Thell) were analyzed for combining ability for quantitative and quality traits. The results indicated significant differences among the parents for general combining ability (gca) and crosses for specific combining ability (sca) for all the characters studied. The gca and sca components of variance were significant for all the traits. However, the gca component of variance was predominant indicating the predominance of additive gene effects for the traits studied. Among the parents Durgapura 65, HD 2285, Lok-1, Raj 1972 and HD 2329 were the best general combiners for grain yield and average to high combiners for tillers per plant, grain yield per spike, grains per spike and 1000-grain weight. The best specific crosses for grain yield were Sonalika x WH 157, HD 2428 x Durgapura 65, Durgapura 65 x Sonalika, HD 2428 x Lok-1 and CPAN 3004 x Raj 1972. The parent Raj 1972, Lok-1 and HD 2285 were the best general combiners for grain yield and protein content, however, Raj 3077 was the best general combiner for protein content. The most suitable specific crosses for protein content were HD 2329 x HD 2285, HD 2428 x Raj 1972 and CPAN 3004 x WH 157. Most of the specific crosses for grain yield as well as protein content involved high x average, average x average and average x poor general combiners. To ensure further increase in grain yield along with high protein, combinations of desirable yield components is advocated. Inclusion of F(1) hybrids showing high sca and having parents with good gca, into multiple crosses and/or bi-parental mating, or diallel selective mating could prove a worthwhile approach for further improvement of grain yield in bread wheat.     

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.     

Diallel analysis to predict heterosis and combining ability for grain yield,yield components and bread-making quality in bread wheat (T. aestivum)

Combining ability for grain yield, yield components, and several agronomic and qualitative traits, was studied in a seven-parent diallel cross. The 21 F₁ hybrids and the seven parental cultivars were grown in replicated plot trials sown at normal seed density in three locations in the years 1992 and 1993. The effects of general combining ability (gca) were highly significant for all the traits measured with the exception of seeds per spikelet, while the specific combining ability (sca) effects were statistically significant for grain yield, plant height, heading time, for all the yield components, and for the Chopin alveographic parameters P and P/L ratio. For the majority of the traits measured gca was greater than sca. Standard heterosis (sh) for grain yield, i.e., the superiority of the hybrids over the best pure line cultivar(cv Eridano), was only 3.3%, confirming previous finding which indicate sh effects in the range of 10%. The most interesting hybrid derived from the cross Maestra x Golia revealed a yield level approaching that of the highest yielding cv Eridano but appeared more interesting because of its reduced plant height and superior bread-making quality, signifying a selling price 30% higher. It was concluded, therefore, that the first generation of hybrids, likely to appear on the market in the next few years, will be characterized by a yield potential only slightly superior to that of the best standard cvs but associated with other desirable traits, such as bread-making quality.     

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.     

Genome‐wide association analyses reveal the genetic basis of combining ability in rice

Combining ability is a measure for selecting elite parents and predicting hybrid performance in plant breeding. However, the genetic basis of combining ability remains unclear and a global view of combining ability from diverse mating designs is lacking. We developed a North Carolina II (NCII) population of 96 Oryza sativa and four male sterile lines to identify parents of greatest value for hybrid rice production. Statistical analyses indicated that general combining ability (GCA) and specific combining ability (SCA) contributed variously to different agronomic traits. In a genome‐wide association study (GWAS) of agronomic traits, GCA and SCA, we identified 34 significant associations (P < 2.39 × 10^?7). The superior alleles of GCA loci (Ghd8, GS3 and qSSR4) accumulated in parental lines with high GCA and explained 30.03% of GCA variance in grain yield, indicating that molecular breeding of high GCA parental lines is feasible. The distinct distributions of these QTLs contributed to the differentiation of parental GCA in subpopulations. GWAS of SCA identified 12 more loci that showed dominance on corresponding agronomic traits. We conclude that the accumulation of superior GCA and SCA alleles is an important contributor to heterosis and QTLs that greatly contributed to combining ability in our study would accelerate the identification of elite inbred lines and breeding of super hybrids.     

几个优良籼稻亲本品质性状的配合力和杂种优势分析

以3个不育系和10个恢复系为材料,采用NCII交配设计研究10个米质性状的配合力和杂种优势。结果表明：①大多数品质性状的量值介于双亲之间,除粒重表现一定的超亲优势、垩白度和粒宽表现一定的正向平均优势外,其他品质性状优势不明显。②杂种稻米的品质性状主要受不育系或恢复系的影响,其中粒长、粒宽和直链淀粉含量3个性状,不育系的影响要高于恢复系;而对于整精米率、粒重、垩白率、垩白度和糊化温度,则恢复系的影响要高于不育系。③就优质育种的利用价值而言,不育系以广占63-4S为好,恢复系以扬稻6号为好,R527、镇恢084次之,用上述亲本选配的杂交组合米质较好;恢复系特青、盐恢559表现为一般配合力效应低,特殊配合力方差小,优质育种利用价值不大。     

Genetics of transgressive segregation for yield and yield components in wheat

Three crosses of spring wheat (Triticum aestivum L. em Thell) involving six cultivars (WC29, WH291, SGP 14, RAJ. 1972, WH377 and HD 2329) were selected on the basis of combining ability analysis to study genetics of transgressive segregation for tillers/plant, grains/spike, 1000 grain weight and grain yield/plant using various mating designs. Diallel analysis indicated that both additive and non- additive components were significant for all the characters. On the basis of general combining ability and specific combining ability effects, the parents WH 291 and WH 377 were found to be good general combiners for tillers/ plant, 1000 grain weight and grain yield/plant. For grains/spike SGP 14 was found to be a good general combiner. The cross WH 377 × HD 2329 for tillers/plant, SGP 14 × Raj. 1972 for grains/spike and grain yield/plant and WC 29 × WH 291 for 1000 grain weight were found to be good cross combinations. Generation mean analysis indicated that the additive-dominance model was inadequate for all the characters in all the crosses except for 1000 grain weight in WC29 × WH291. Additive component was more pronounced than non-additive components for all the characters in all the crosses except for tillers/ plant in WH 377 × HD 2329. Predictions for transgressive segregants from F₃ was more accurate than that from generation mean analysis. However, prediction from both the sources were equally efficient if additive-dominance model was adequate. In general, observed frequencies of transgressive segregants were more in F₂ and BIPi than F₄ but the majority of them were discarded on progeny testing. Biparental mating had an impact in increasing the frequencies of transgressive segregants for different characters in all the crosses. The crosses, WH 377 × HD 2329 for grain yield/ plant, SGP 14 × Raj. 1972 for tillers/plant and WC 29 × WH 291 for grains/spike and 1000 grain weight were found to be potential crosses for transgressive segregants. A comparison of combining ability of parents and crosses, and observed and predicted frequencies of transgressive segregants indicated that the potential crosses for transgressive segregants were those that had high sea effects and involved high and low general combiners. The crosses involving low general combiners irrespective of their sea effects showed poor performance with respect to transgressive segregation.     

杂交水稻主要农艺性状配合力遗传力分析

选用生产上常用的5个杂交水稻不育胞质与5个高配合力恢复系,采用NCII模式对产量性状、芒、粒形等主要农艺性状的亲本配合力、方差贡献率及遗传力进行分析。结果表明：胞质效应和恢复系效应在所测性状中均达到显著或极显著差异水平,芒长、粒形、单穗重受环境的影响较小;产量和收获指数的非加性遗传作用明显,受环境效应的影响也较大。亲本效应方面,恢复系的效应远大于胞质效应;产量性状JW型（爪哇型）胞质和蜀恢527的一般配合力效应较高,同时JW型胞质和R21属于Ⅰ类亲本;粒形性状G型（冈型）胞质和蜀恢527的一般配合力效应最高,同时均属于Ⅰ类亲本。     

Assessment of cytoplasmic differences of near-isonuclear male-sterile lines in pearl millet

Four near-isonuclear polycytoplasmic versions of 81A and two of Pb 402A male-sterile lines of pearl millet (Pennisetum typhoides) were used in factorial matings with five inbred male testers in different combinations in three sets. The cytoplasmic differences were studied for several agronomic traits using mean values and general combining effects (gca) of male-sterile lines, and specific combining ability effects of hybrids. The fertility/ sterility behaviour of different male-sterile lines in crosses with common male parents was also studied. Significant differences among near-isonuclear polycytoplasmic lines were observed in mean values for a few traits such as plant height, leaf length and peduncle length, but the differences for combining ability were more pronounced. The A₃ cytoplasm was a better general combiner than the A₂ cytoplasm for grain yield and both A₂ and A₃ cytoplasms were better general combiners for leaf length and peduncle length. In addition, superiority of A₃ cytoplasm for gca was observed for plant height and ear characters over the A₂ cytoplasm in set II. A differential behaviour of cytoplasms, both in combination with a common pollinator and across pollinators, was observed for several traits. The results provide evidence for the distinctiveness of different cytoplasmic sources in pearl millet and for the influence of cytoplasmic factors on the phenotypic expression of nuclear genes. A diversification of male sterility sources in the breeding of pearl millet hybrids is suggested.     

Heterosis and components of genetic variation for protein and lysine content in some grain sorghums

Summary Twelve varieties representing six geographical regions and nine taxonomic groups from a World Collection ofSorghum were used in a diallel and line X tester analysis of the nature of genetic variation for protein and lysine content. In a majority of crosses, heterosis was negative for protein, and positive for lysine.Analysis of combining ability indicated that both additive and non-additive variation were important for protein, while the non-additive component was predominant for lysine. Heterosis inlow Xlow andmedium Xlow crosses also indicated the presence of substantial non-allelic gene interactions for both the characters.Associations between protein, lysine and yield in parents were significantly different from those in the hybrids indicating considerable scope for genetic manipulation. Negative correlations between protein and lysine (% protein) and between protein and grain yield were very low. The results indicated that a high level of protein and moderately high lysine may be incorporated in high yielding varieties of sorghum, by using Caudatum kaura, Roxburghii shallu and durra from Nigeria and Sudan in the hybridisation programme.     

Parental Selection of Hybrid Breeding Based On Maternal and Paternal Inheritance of Traits in Rapeseed (Brassica napus L.)

Parental selection is crucial for hybrid breeding, but the methods available for such a selection are not very effective. In this study, a 6×6 incomplete diallel cross was designed using 12 rapeseed germplasms, and a total of 36 hybrids together with their parental lines were planted in 4 environments. Four yield-related traits and seed oil content (OC) were evaluated. Genetic distance (GD) was estimated with 359 simple sequence repeats (SSRs) markers. Heterosis levels, general combining ability (GCA) and specific combining ability (SCA) were evaluated. GD was found to have a significant correlation with better-parent heterosis (BPH) of thousand seed weight (TSW), SCA of seeds per silique (SS), TSW, and seed yield per plant (SY), while SCA showed a statistically significant correlation with heterosis levels of all traits at 1% significance level. Statistically significant correlations were also observed between GCA of maternal or paternal parents and heterosis levels of different traits except for SS. Interestingly, maternal (TSW, SS, and OC) and paternal (siliques per plant (SP) and SY) inheritance of traits was detected using contribution ratio of maternal and paternal GCA variance as well as correlations between GCA and heterosis levels. Phenotype and heterosis levels of all the traits except TSW of hybrids were significantly correlated with the average performance of parents. The correlations between SS and SP, SP and OC, and SY and OC were statistically significant in hybrids but not in parents. Potential applications of parental selection in hybrid breeding were discussed.     

抗除草剂杂交籼稻亲本的配合力分析

本试验以6份新育成的抗除草剂籼型恢复系为父本,5份生产上广泛应用的不育系为母本,采用不完全双列杂交设计配制了30份杂交组合,对其苗期除草剂抗性和主要农艺性状配合力进行了分析。除草剂抗性鉴定表明,亲本恢复系及三系杂交组合抗性接近完全,两系杂交组合抗性达90%以上。配合力分析表明,不育系除单株有效穗数外其他农艺性状的一般配合力均达到极显著差异;恢复系间一般配合力在所有性状中均达到显著或极显著差异;杂交组合间特殊配合力方差仅在单株产量、结实率、播始历期和千粒重4个性状中达到显著或极显著差异。不育系中,金科1A在单株产量、结实率等7个性状上的一般配合力均为最高,但其特殊配合力方差最小;广占63-4S在千粒重性状上具有最高的一般配合力、最大的特殊配合力方差,在播始历期上具有最高的一般配合力负效应;C815S在株高上的一般配合力负效应最大,同时特殊配合力方差较高。恢复系中,华抗恢101在单株有效穗数上具有最高的一般配合力和特殊配合力方差;华抗恢104在穗长上具有最高的一般配合力,在播始历期上具有最高的一般配合力负效应;华抗恢105在单株产量、结实率等性状上具有最高的一般配合力,在株高上具有最高的一般配合力负效应;华抗恢106在千粒重性状上具有最高的一般配合力和特殊配合力效应方差。利用抗除草剂恢复系配制杂交组合,不仅可以改良其除草剂抗性,也可以通过广泛测配,选择一般配合力强、特殊配合力方差大的亲本配组育成强优势组合。     

Somatic embryogenesis in Zea mays L.

Summary Combining ability studies with respect to such green fodder quality characteristics as oxalic acid, calcium, sodium, potassium and green fodder yield were carried out in a 12 × 12 diallel cross set in pearl millet (Pennisetum typhoides (Burm) S. & H.). With regard to differential expression of gene effects, studies for quality traits were carried out in different seasons and on different plant parts. The relative proportions of general and specific combining variances indicated the preponderance of non-additive genetic variance. Parents possessing desirable fodder quality characteristics were identified on the basis of combining ability and per se performance, and selection criterion for crosses was discussed. It was recommended that leaf portion should be biochemically analysed and manipulated in an environment when the genes are expressed.Part of the Ph. D. dissertation submitted to the Punjab Agricultural University by the senior author in partial fulfilment of the requirements for the degree     

高粱穗部主要性状的配合力分析

以4个高粱不育系13163A、1358A、128A和407A,以及6个恢复系9.1R、213R、272R、381R、矮182R和早21R为试验材料,按照不完全双列杂交设计(NCII),对其F1的穗部主要性状进行配合力分析。结果表明:亲本穗部主要性状存在显著的遗传差异,主要表现为加性基因效应遗传的性状有:穗长、一级枝梗数、二级枝梗数、穗粒数;狭义遗传力大小顺序分别为:二级枝梗数一级枝梗数穗长穗粒数穗粒重千粒重。不同亲本的一般配合力(GCA)和特殊配合力(SCA)在不同穗部性状间存在较大差异。不育系407A和恢复系早21R、9.1R是综合性状较好的亲本材料,利用它们可组配出产量较高的杂交组合。恢复系272R组配的杂交种具有穗粒数较多、千粒重较小的特点,能满足市场上对小粒高粱的需求。深入分析高粱杂交亲本穗部主要性状表现,有利于对亲本材料的进一步了解和利用。     

小麦品种间杂种优势与配合力分析

对９个冬小麦亲本１３个性状的杂种优势分析表明,产量性状的杂种优势较强,品质性状的杂种优势较弱。籽粒蛋白质含量、硬度和湿面筋、干面筋含量偏低亲或中亲遗传,沉淀值偏高遗传,存在较明显杂种优势（１６．１－２３．８％）。     

小麦品质和农艺性状的配合力分析

依据田间试验和室内生化分析数据,按ｐｘｑ交配模式,对１０个冬小麦亲本２５个杂交组合Ｆ１的蛋白质含量,赖氨酸含量,穗粒重和千粒重的一般配合力和特殊配合力效应进行了分析。结果表明,除蛋白质含量的ｓｃａ外,其余性状的ｇｃａ和ｓｃａ效应均达极显著水平,表明在本研究中,基因的加性效应和非加性效应均起重要作用。由于一般配合力差δ＾２ｓｃａ大,表明基因的加性效应更加重要。根据ｇｃａ,ｓｃａ和δ＾２ｓｃａ估值,分     

Prediction of single-cross hybrid performance for grain yield and grain dry matter content in maize using AFLP markers associated with QTL

Prediction methods to identify single-cross hybrids with superior yield performance have the potential to greatly improve the efficiency of commercial maize (Zea mays L.) hybrid breeding programs. Our objectives were to (1) identify marker loci associated with quantitative trait loci for hybrid performance or specific combining ability (SCA) in maize, (2) compare hybrid performance prediction by genotypic value estimates with that based on general combining ability (GCA) estimates, and (3) investigate a newly proposed combination of the GCA model with SCA predictions from genotypic value estimates. A total of 270 hybrids was evaluated for grain yield and grain dry matter content in four Dent × Flint factorial mating experiments, their parental inbred lines were genotyped with 20 AFLP primer-enzyme combinations. Markers associated significantly with hybrid performance and SCA were identified, genotypic values and SCA effects were estimated, and four hybrid performance prediction approaches were evaluated. For grain yield, between 38 and 98 significant markers were identified for hybrid performance and between zero and five for SCA. Estimates of prediction efficiency (R ²) ranged from 0.46 to 0.86 for grain yield and from 0.59 to 0.96 for grain dry matter content. Models enhancing the GCA approach with SCA estimates resulted in the highest prediction efficiency if the SCA to GCA ratio was high. We conclude that it is advantageous for prediction of single-cross hybrids to enhance a GCA-based model with SCA effects estimated from molecular marker data, if SCA variances are of similar or larger importance as GCA variances.     

Genetic parameters of intra- and inter-specific hybrids of Eucalyptus globulus and E. nitens

Intra-specific hybrids within Eucalyptus nitens and E. globulus were compared directly with inter-specific E. nitens × globulus using common parents. Diameter (age 2, 4, 6, 10 years) and Pilodyn (age 6 years) were used as indirect measures of growth and wood density, respectively. Genetic parameters were estimated for all cross types and traits. A direct comparison of the general combining ability with the general hybridising ability was made. Heterosis was estimated for intra- and inter-specific hybrids. The intra-specific hybrids in both species exhibit intermediate heritabilities and levels of additive genetic variance. The inter-specific F1 E. nitens × globulus exhibited high incidence of abnormalities at young ages and high levels of later age mortality. The mean performance of surviving inter-specific F1 hybrids was generally intermediate for all traits, to varying degrees, between the parental intra-specific crosses, and there is little evidence for significant heterosis. Different provenances of E. globulus may exhibit different responses to hybridisation. In this case, the inter-specific F1 hybrids, using Taranna E. globulus, appear to perform worse on average than those using King Island provenance for growth traits. There is evidence that specific hybrid families are produced that outperform most of the pure species families for one or other of the traits examined; however, there is no reliable quantitative genetic method of predicting which parents should be used.     

Genetic variation in grain-filling ability in dwarf pearl millet [Pennisetum glaucum (L.) R. Br.] restorer lines

The d2 dwarfing gene in pearl millet [Pennisetum glaucum (L.) R. Br.] carries a yield penalty due to an associated reduction in individual grain mass. This reduction, however, varies with genetic background, indicating that it may be possible to select against poor grain filling in d2 dwarfbackgrounds, given an effective measure of grain filling. This study was conducted to assess genetic variability forgrain-filling ability (in contrast to simply grain size),and its relationship to grain yield,indwarf pearl milletrestorer (R) lines. The grain-filling ability (GFA) of an individual R line was defined as the least squares estimate of its effect on individual grain mass in the analysis of variance, following a linear covariance adjustment for grain number. The study was based on 93dwarf hybrids involving31 d2 dwarfR-lines, evaluated over 3 years. Half of the variation in individual grain mass in the 93 hybrids was related to variation in grain number. Covariance adjustment in individual grain mass for grain number resulted in highly significant differences among hybrids and R lines in GFA. The R-line combining ability for GFA accounted for 26% of the variation in the R-line combining ability for yield, compared to 46% for the combining ability for grain number, and just 8% for the combining ability of individual grain mass. The combining ability for GFA was independent of the combining ability for various pre-flowering effects, including grain number, but was related to the combining ability for individual grain mass and harvest index. Improvement in individual grain mass achieved through selection for GFA should translate directly into yield improvement, whereasimprovement by direct selectionfor individual grain mass is less-likely to do so. Received: 9 April 2000 / Accepted: 16 May 2000     

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.