Application of the concepts of test value and varietal value to the study of genetic advance in recurrent selection: A synthesis

Summary The concepts of varietal ability and value in test of a genotype or group of genotypes are specified according to the type of variety (clones, hybrids, lines, synthetics) and the testing system (phenotype, S₁, General combining ability (GCA), line value, General synthesizing ability (GSA)). Varietal ability and value in test are to be considered as any quantitative characters for which it is possible to define additive effects, dominance effects and epistasis. For diploids in recurrent selection, only additive effects and additive× additive epistasis for varietal value contribute to genetic advance. Genetic advance depends on the covariance between the testing value of the parents and the test value of the progeny by intercrossing. Such a parent-offspring covariance is, in the absence of epistasis and inbreeding, one half of the covariance between additive effects in test of the parents and additive effects in test of their progenies. This allows one to express directly the genetic advance in varietal value according to the type of variety and to the testing system used in recurrent selection. The case of biallelic populations is considered; it allows some discussions on the relative values of components of variance of varietal and test values.     

Reciprocal recurrent selection applied to Coffea canephora Pierre. III. Genetic gains and results of first cycle intergroup crosses

Breeding of the allogamous Coffea canephora species in Côte d'Ivoire is based, from 1984 onward, on a reciprocal recurrent selection scheme exploiting the good performance of crosses between two genetically distinct populations (‘Guinean’ and ‘Congolese’). Results are presented here on intergroup cross progenies evaluated on individual plant basis over a six year period in three fully randomized trials planted in 1985 and 1986. Yields of the bi-parental crosses varied from 16 to 1407% of the mean of the two commercial clonal cultivars and the most productive crosses were also the most vigourous. Selection of individual trees and cross progenies, based on a combined individual-family index, was carried out for high yield, good vigour of young plants and moderate canopy diameter of adult trees. Expected genetic gains, as estimated in relation to the mean of all crosses, for selection of 5% of the best trees, were high for yield (over 60%), moderate for young plant vigour (14 to 18%), and, as intended, low for canopy diameter. In addition, selection by independent culling for leaf rust resistance and bean weight was carried out. As a result, 6 promising crosses with yield similar or higher than those of the control cultivars, were selected as candidates for commercial use, as well as 65 individual trees, within 19 different crosses. Expected genetic gain for yield for these selected trees is about 60% compared to the average yield of the most productive control clone. Continuation of the programme is discussed, with specific attention to further progress expected for quality traits.     

籼型杂交水稻农艺性状的配合力研究

如何有效利用杂种优势已成为水稻增产的关键。本研究按照NCII遗传交配设计,将三系野败型杂交水稻的恢复系和微核心种质构成的115份优异籼稻品种,分别与4个两系不育系及1个三系不育系测交,分析各农艺性状配合力、遗传力及相互关系。结果表明,除单株有效穗数、主穗实粒数外,其他农艺性状一般配合力差异均达到极显著水平;除单株有效穗数外,其他各农艺性状特殊配合力差异也均达到极显著水平。同一组合的不同性状、同一亲本的不同组合所表现出的特殊配合力效应都有所不同,表明亲本的一般配合力水平与特殊配合力间没有固定的联系。在育种实践中,选取一般配合力高的亲本,同时兼具较高特殊配合力是获取高产杂交稻组合的关键。     

Comparison of two crossing and four selection schemes for yield, yield traits, and slow rusting resistance to leaf rust in wheat

The most important breeding objectives in crop improvement are improving grain yield, grain quality, and resistances to various biotic and abiotic stresses. The objectives of our study were to compare two crossing and four selection schemes for grain yield, yield traits, and slow rusting resistance to leaf rust (Puccinia recondita) based on additive genes in wheat (Triticum aestivum), and to identify the most efficient crossing and selection methodologies in terms of genetic gains and cost efficiency. Segregating populations were derived from 18 simple crosses and the same number of top (three-way) crosses. Half of the crosses were derived from Yecora 70 and the other half from Veery #10 as the common leaf rust susceptible parents. The four selection schemes were: pedigree, modified bulk (F₂ and F₁-top as pedigree, selected lines in F₃, F₄, F₂-top, F₃-top as bulk; and pedigree in F₅ and F₄-top populations), selected bulk (selected plants in F₂, F₃, F₄, F₁-top, F₂-top and F₃-top as bulk; and pedigree in F₅ and F₄-top populations), and nonselected bulk (bulk in F₂, F₃, F₄, F₁-top, F₂-top and F₃-top; and pedigree in F₅ and F₄-top populations). A total of 320 progeny lines, parents and checks were tested for grain yield, other agronomic traits and leaf rust resistance during the 1992/93 and 1993/94 seasons in Ciudad Obregon (Sonora State, Mexico) which represents a typical high yielding irrigated site. The influence of the type of cross and the selection scheme on the mean grain yield and other traits of the progenies was minimal. The selection of parents was the most important feature in imparting yield potential and other favourable agronomic traits. Moreover, the highest yielding lines were distributed equally. Progeny lines derived from Veery #10 crosses had significantly higher mean grain yield compared to those derived from the Yecora 70 crosses. Furthermore, a large proportion of the highest yielding lines also originated from Veery #10 crosses. Mean leaf rust severity of the top cross progenies was lower than that of the simple cross progenies possibly because two parents contributed resistance to top cross progenies. Mean leaf rust severity of the nonselected bulk derivatives was twice that of lines derived from the other three schemes. Selected bulk appears to be the most attractive selection scheme in terms of genetic gains and cost efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

大豆亲本品种数量性状的配合力分析

采用6个大豆亲本双列杂交设计,研究亲本及F_1、F_2的配合力.各性状的一般和特殊配合力方差分析多达显著水平.一般配合力效应表现较好的有豫豆6号和郑8431.在F_1和F_2中同一性状不同组合或同一组合不同性状间的特殊配合力效应具有显著差异.在不同世代中根据遗传力的大小,应对不同性状进行选择.单株粒重遗传力较低,不易在早期世代选择.     

Combining ability and heterosis under pest epidemics in a broad-based global wheat-breeding population

Wheat breeders rarely apply population improvement schemes or select parental sources according to combining ability and heterotic patterns. They rely on pedigree selection methods for breeding new cultivars. This experiment was undertaken to assess the advantages of using diallel crosses to define combining ability and understand heterosis in a broad‐based wheat‐breeding population across different environments affected by yellow rust. Sixty‐four genotypes derived from a full diallel mating scheme were assessed for grain yield in two contrasting growing seasons at two locations for two consecutive years. Parental genotypes showed significant combining ability for grain yield that was affected by yellow rust and genotype‐by‐environment (GE) interactions, both of which affected heterosis for grain yield. Significant GE interactions suggested that decentralized selection for specific environments could maximize the use of this wheat germplasm. Cultivar effects and specific heterosis were the most important factors influencing grain yield. Some crosses capitalized on additive genetic variation for grain yield. This research shows the power of available quantitative breeding tools to help breeders choose parental sources in a population improvement programme.     

Heritability and correlations among traits in four-way soybean crosses

Recurrent selection programs can use several crossing arrangements to synthesize basic populations for breeding purposes. This study evaluated heritability and correlations among traits in forty-five populations of F_2[4] and F_2:3[4] generations obtained throughout four-way crosses [4] between two dialells among ten semi-late and ten late maturity soybean (Glycine max L. Merrill) genotypes. The generation advance was made by the TSHD (Thinned Single Hill Descent) method and all individual plants were evaluated for the following traits: number of days to flowering (NDF) and maturity (NDM), plant height at flowering (PHF) and maturity (PHM), agronomic value (AV) and seed yield (YLD). Four-way crosses associated with TSHD method provided high genetic variability in the populations with little or no reduction from F_2[4] to F_2:3[4]. The ‘narrow sense’ heritability estimated using correlations between F_2[4] and F_2:3[4] plants was lower than but closely related to broad-sense estimates. The populations 21(IAC-4 × IAC-9) × (GO79-1039 × Paranagoiana) and 23 (IAC-4 × Santa Rosa) × (GO79-1039 × Tropical) were the most productive with high variability in F_2[4] and/or F_2:3[4] generations. Genotypic correlations measured were generally high, positive and consistent for the two generations except for those related to AV and YLD. These results indicate that the use of four-way crosses can be an efficient method to increase the genetic base of populations for recurrent selection or cultivar development. Significant AV × YLD correlations indicated visual selection can be used as additional selection criteria for improving populations. This revised version was published online in August 2006 with corrections to the Cover Date.     

三交和双交组合加性-显性-母体效应的遗传模型及蒙特卡罗模拟分析

本文根据广义遗传模型的建模原理,按照三交和双文组合方式的交配设计,分别提出了分析 作物三交和双交组合数量性状的加性－显性－母体效应的遗传模型（ＡＤＭ模型）,给出了不同环境下各 世代群体的遗传效应分量。应用ＭＩＮＱＵＥ（１）法对包括不同世代的三交和双交组合的ＡＤＭ模型进行 了蒙特卡罗模拟分析,结果表明：采用亲本和三（双）交Ｆ_１两个世代分析三（双）交组合的ＡＤＭ模型 时,即可获得方差分量的无偏估计值和遗传效应的无偏预测值;采用包括单交Ｆ_１在内的三个世代联合 分析时,更有利于提高显性等遗传效应方差分量的估算效益。最后讨论了利用单交组合预测三交和双 交组合遗传表现的有关问题。     

三交和双交组合加性-显性-母体效应的遗传模型及蒙特卡罗模拟分析

本文根据广义遗传模型的建模原理, 按照三交和双交组合方式的交配设计, 分别提出了分析作物三交和双交组合数量性状的加性-显性-母体效应的遗传模型(ADM模型), 给出了不同环境下各世代群体的遗传效应分量。 应用MINQUE(1)法对包括不同世代的三交和双交组合的ADM模型进行了蒙特卡罗模拟分析, 结果表明： 采用亲本和三(双)     

Heterosis: its genetic basis and its utilisation in plant breeding

Summary To discuss the utilisation of heterosis in plant breeding it is necessary to consider its genetic basis. In cross fertilized plants in which the phenomenon of heterosis is very important, such as maize, it appears that a great part can be explained by the mutation load. However in both autogamous or allogamous plants it is difficult to exclude a role of marginal overdominance which could justify hybrid variety production.To know in a relatively short time whether it will be better to develop lines or hybrids it is necessary to know the parameters (mean and variance) of the distribution of all varieties of a given type which can be derived from the breeding population. A numerical application allows the determination of the situations where the best single crosses will be better than the best lines. With some approximate values of the necessary parameters, the case of grain yield in maize and in wheat is briefly discussed.To develop lines or hybrids, a general strategy is given. Its main axis is population improvement which has to be adapted to the type of varieties to develop. It is shown that according to the situation the type of varieties to develop can change with time: single crosses can be more justified at the beginning and lines can be justified later if all heterosis were fixable. To develop hybrids the best scheme is reciprocal recurrent selection with half-sib or full-sib progenies followed by pedigree reciprocal selection initiated by full-sib progeny tests. With this strategy there is no problem of the prediction of hybrid performances. However it seems possible to develop some predictors of heterosis or of specific combining ability between two lines using some criteria of genetic distances. Preliminary results with distances computed from the behaviour of lines in a two-tester top cross design appears very stimulating.Communication given at the206th meeting of teh Genetical Society of Grain Britain     

玉米三交种郑三3号高产原因分析

用郑三3号的3个亲本自交系,组配成6个正反单交组合和6个以单交为母本的三交组合,进行产量比较和性状调查。结果表明,6个单交组合之间的产量差异不显著,6个三交组合之间的产量差异亦不显著,说明这3个自交系相互间的配合力都很高,这是郑三3号的产量达到了优良单交种产量水平的内在因素。     

Gene action in the resistance of peppers (Capsicum annuum) to Phytophthora stem blight (Phytophthora capsici L.)

Summary The variation among single and three-way crosses derived from seven lines of pepper (Capsicum annuum) selected for resistance to Phytophthora capsici was analyzed into statistical-genetic components due to general and specific combining abilities and additive-by-additive epistasis. Indirect evidence of higher order epistasis was also estimated. Epistasis was a principal source of variation. It was correlated with the level of aggressiveness of the pathogen. A breeding method such as recurrent selection seems to be the most appropriate to improve resistance to the fungus.     

Genetic divergence for cross prediction in potato

To study the effectiveness of genetic divergence for cross prediction in potato, progeny means, heterosis and specific combining ability effects were correlated with parents’ genetic distances (D values) estimated under six in vitro and four in vivo conditions, for tuber yield in 72 crosses (18 × 4) of 22 parents under autumn crop conditions for three successive generations. Genetic distances under in vitro conditions had no relationship with the progeny means for tuber yield. Whereas, those under in vivo conditions in the autumn seasons were positively associated with the progeny means. Similarly, heterosis for tuber yield had a stronger relationship with genetic distances based on an in vivo crop than those based on an in vitro crop. All correlation coefficients between genetic distances and specific combining ability effects were non-significant. The magnitudes of the significant correlation coefficients showed that genetic divergence can be used as an indirect parameter of moderate effectiveness in selecting parents to produce heterotic high yielding progenies. Such cross predictions, however, would be effective only if parents are evaluated under the conditions similar to those under which crosses are likely to be evaluated. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding for yield in dry bean (Phaseolus vulgaris L.)

Strategies employed by dry bean breeders to improve yield include early generation testing, ideotype breeding, selection for physiological efficiency, and selection based on genotypic performance and combining ability across gene pools of Phaseolus vulgaris. Ideotype breeding has been successfully deployed to improve yield in navy, pinto and great northern seed types. The ideotype method is based on an ideal plant architecture to which breeders target their selection. Breeding for physiological efficiency is important in combining increased biomass, high growth rates and efficient partitioning. Genotypic performance and combining ability are also critical for yield improvement, since crosses between gene pools can exhibit negative combining ability and problems with lethality, whereas interracial crosses within the same gene pool exhibit the greatest potential. Breeders must work within specific constraints for growth habit, maturity, seed quality and disease resistance. A three-tiered pyramidal breeding strategy is proposed to facilitate yield improvement in dry bean. Breeding of elite, agronomically acceptable germplasm within the same market class is restricted to the apex of the pyramid. The intermediate level has fewer constraints and greater access to diverse germplasm. Interracial crosses within the same gene pool are utilized to exploit genetic differences within adapted material. Extracting genetic diversity from unadapted sources, including wild germplasm and other Phaseolus species, is conducted at the base of the pyramid. The objective of this breeding strategy is the movement of improved germplasm towards the apex, using different breeding procedures to optimize improvement at each tier of the breeding pyramid. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic analysis of postharvest flower longevity in Asiatic hybrid lilies

To investigate the genetic regulation of postharvest flower longevity in Asiatic hybrid lilies (Lilium L.), 10 cultivars and 45 progenies were forced, harvested and evaluated under standardised conditions in growth chambers. Analysis of variance for individual flower longevity indicated highly significant (p < 0.001) variation among parents, among progenies and among descendants within progenies. High broad-sense heritability (0.79) calculated at the individual plant level indicated that selection for long individual flower longevity can be expected to be very effective. General combining ability (GCA) effects were highly significant ( p < 0.001), and the estimated narrow-sense heritability was high (0.74). Therefore, individual flower longevity of a genotype can be used as an indication for its breeding value. Although deviating results can be expected as specific combining ability (SCA) effects were also significant ( p = 0.046). Small, but significant correlations between individual flower longevity and other plant characters were found. The impact of these correlations on the selection efficiency for improved postharvest performance of lily inflorescences is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance to Coffee Berry Disease in Arabica Coffee Variety 'Ruiru 11'

General and specific combining ability effects for resistance to coffee berry disease (Colletotrichum coffeanum Noack sensu Hindorf) in Coffea arabica variety ‘Ruiru 11’ and its parents were estimated according to the North Carolina Design II. A set of nine male parents and eleven females were crossed in all possible combinations to generate ninety-nine single cross hybrids constituting the ‘Ruiru 11’ variety. The parents and their hybrid progenies were screened and the mean score data analyzed for possible genetic variation. The contributions of the parents to the total genetic variance were low hence the low narrow sense heritability (h²= 0.04). The male and female parents accounted for 9.36 % and 0.96 %, respectively, while their interaction effect accounted for 89.68 %. In the analysis of combining ability for resistance, ExB3.96 and Cat.90 exhibited significant negative general combining ability (GCA). Specific combining abilities (SCA) were negative and significant for Cat.86 × ExB3.99, Cat.127 × ExB3.116 and Cat.119 × ExB3.879 crosses. The implications of these results in selection for highly resistant ‘Ruiru 11’ progenies are discussed.     

Selection for combining ability in Pyrethrum,Chrysanthemum cinerariaefolium Vis.

Summary Pyrethrum is grown for the insecticidal pyrethrins, present mainly in the flowers. It is a cross-fertilizing perennial which can be propagated vegetatively. The main selection criteria are fresh flower yield and pyrethrins content; important secondary criteria are flower size and resistance to lodging; dry matter content varies very little.For these traits, the breeding values (general combining ability) of 22 clones were determined by means of a polycross. The heritabilities were estimated from the regressions of the polycross progeny data on those of the female parent clones; these were: flower yield 0.31; pyrethrins content 0.70; flower size 0.80 and resistance to lodging 0.87. For dry matter content no correlation between parents and progenies could be measured. Independent heritability estimates, obtained from 33 single crosses between clones not represented in the polycross, were: flower yield 0.56; pyrethrins content 0.64. Of 18 single crosses, made between clones also represented in the polycross, the actual flower yields and pyrethrins contents were highly correlated with the estimated yields and contents which were calculated from the breeding values of the parent clones (r = +0.87 and +0.86, respectively).These data show that the performance of single crosses is caused predominantly by the general combining ability of the parent clones. Therefore, single crosses offer no fundamental advantages over synthetic varieties, so that, for practical reasons, the latter are preferred. At the same time it is recommended to practise reciprocal recurrent selection as this will exploit the general combining ability in the initial stages of selection and the specific component increasingly in later selection cycles.Published with the approval of the Director of Agriculture, Kenya.     

Diallel analysis of tolerance to aluminium in alfalfa

Summary Acid soils having high levels of aluminium (Al) can drastically reduce yields in alfalfa and the most economically viable solution to the problem appears to be the development of Al-tolerant cultivars. To assist with the choice of a breeding method, a six-parent alfalfa diallel (crosses and reciprocals included but not parents) was evaluated in Al-toxic nutrient solution in terms of height (HT) and dry weight (DW). General combining ability was significant for both traits and constituted the majority of the genetic variation. Specific combining ability was significant only for HT and reciprocal effects were significant only for DW. Tolerance appeared to be at least partially dominant to sensitivity. Results indicate that a mass selection scheme, such as recurrent phenotypic selection, may be effective in increasing levels of tolerance in at least some alfalfa populations and that minor grains may also be achieved through exploiting non-additive genetic variation.     

Winter × spring wheat hybridization - a promising avenue for yield enhancement

Introgression of the winter gene pool into spring wheat is being considered as one of the strategies to break through the yield plateau. However, little information is available on the combining ability of these two important but distinct groups of wheats in Indian conditions. Therefore, the present study was undertaken to determine the combining ability and gene action of yield and yield attributes in winter × spring wheat crosses. Seventy F₁ progenies developed by 14 winter and five spring wheat lines using a line × tester design were evaluated, along with their parents, for yield and yield attributes in a randomized complete block design under field conditions. The mean squares for all the characters studied showed highly significant differences. The mean squares due to female × male interactions were significant for all the characters studied except for grains per ear and grain weight per ear. Additive genetic effects were found to play a key role in controlling the expression of days to heading, plant height and spikelets per ear.‘MV 19’ and ‘Stepniak’/‘Karvuna’ among winter and ‘PBW 65’ among spring wheats were good general combiners for most of the yield attributes studied. The estimates for specific combining ability effects suggested that, although general combining ability (GCA) effects of most winter wheats are either average or poor, their combination can give desirable genotypes with spring wheat parents possessing a high GCA.     

混合选择与系谱选择对大豆杂交材料定向选择效果比较的研究

通过多年的系统试验,证明对大豆杂交早期世代的材料,用混合选择法针对成熟期、株高、结莢习性,以及种粒大小等性状进行定向的选择,效果并不次于系谱选择法,但却简便易行。大豆杂交材料经过数代的定向混合选择,便形成为能适应当地条件与要求的优良生态类型。在这个基础上,再用个体选择法进行选择,能较有保证地选育出高产优质抗性強的优良品系。