Power of the joint segregation analysis method for testing mixed major-gene and polygene inheritance models of quantitative traits

Understanding the genetic architecture of quantitative traits can greatly assist the design of strategies for their manipulation in plant-breeding programs. For a number of traits, genetic variation can be the result of segregation of a few major genes and many polygenes (minor genes). The joint segregation analysis (JSA) is a maximum-likelihood approach for fitting segregation models through the simultaneous use of phenotypic information from multiple generations. Our objective in this paper was to use computer simulation to quantify the power of the JSA method for testing the mixed-inheritance model for quantitative traits when it was applied to the six basic generations: both parents (P₁ and P₂), F₁, F₂, and both backcross generations (B₁ and B₂) derived from crossing the F₁ to each parent. A total of 1968 genetic model-experiment scenarios were considered in the simulation study to quantify the power of the method. Factors that interacted to influence the power of the JSA method to correctly detect genetic models were: (1) whether there were one or two major genes in combination with polygenes, (2) the heritability of the major genes and polygenes, (3) the level of dispersion of the major genes and polygenes between the two parents, and (4) the number of individuals examined in each generation (population size). The greatest levels of power were observed for the genetic models defined with simple inheritance; e.g., the power was greater than 90% for the one major gene model, regardless of the population size and major-gene heritability. Lower levels of power were observed for the genetic models with complex inheritance (major genes and polygenes), low heritability, small population sizes and a large dispersion of favourable genes among the two parents; e.g., the power was less than 5% for the two major-gene model with a heritability value of 0.3 and population sizes of 100 individuals. The JSA methodology was then applied to a previously studied sorghum data-set to investigate the genetic control of the putative drought resistance-trait osmotic adjustment in three crosses. The previous study concluded that there were two major genes segregating for osmotic adjustment in the three crosses. Application of the JSA method resulted in a change in the proposed genetic model. The presence of the two major genes was confirmed with the addition of an unspecified number of polygenes. Received: 18 August 2000 / Accepted: 9 March 2001     

Assessment of genes controlling Area under Disease Progress Curve (AUDPC) for stripe rust (P. Striiformis F. Sp. Tritici) in two wheat (Triticum Aestivum L.) crosses

Genetic effects on controlling stripe rust resistance were determined in two wheat crosses, Bakhiawar-92 × Frontana (cross 1) and Inqilab-91 × Fakhre Sarhad (cross 2) using Area under Disease Progress Curve (AUDPC) as a measure of stripe rust resistance. The resistant and susceptible genotypes for crosses were identified by initial assessment of 45 wheat accessions for stripe rust resistance. Mixed inheritance model was applied to the data analysis of six basic populations P ₁, F ₁, P ₂, B ₁, B ₂, and F ₂ in the crosses. The results indicated that AUDPC in cross 1 was controlled by two major genes with additive-dominance epistatic effect plus polygenes with additive-dominance epistatic effects (model E). Whereas in case of cross 2, it was under the control of two major genes with additive-dominance epistatic effect plus additive-dominant polygenes (model E-1). Additive effect was predominant then all other types of genetic effects suggesting the delay in selection for resistance till maximum positive genes are accumulated in the individuals of subsequent generations. Occurrence of transgressive segregants for susceptibility and resistance indicated the presence of resistance as well as some negative genes for resistance in the parents. The major gene heritability was higher than the polygene heritability in B ₁, B ₂ and F ₂ for the crosses. The major gene as well as the polygene heritability was ranging from 48.99 to 87.12% and 2.26 and 36.80% for the two crosses respectively. The highest phenotypic variations in AUDPC (2504.10 to 5833.14) for segregating progenies (BC ₁, BC ₂ and F ₂) represent that the character was highly influenced by the environment. The article is published in the original.     

粳稻穗部性状遗传分析

从粳稻(Oryza sativa ssp. japonica)RIL群体中选取每穗颖花数极端少的品系丙堡3201和丙堡3205及每穗颖花数极端多的品系丙堡3145和丙堡3214, 配制丙堡3201×丙堡3145和丙堡3214×丙堡3205两个组合的P₁、P₂、F₁、B₁、B₂和F₂ 6个世代, 调查每穗颖花数、每穗实粒数、穗长、一次枝梗数和二次枝梗数的表型分布, 并运用主基因+多基因混合遗传模型,对这5个性状进行了遗传分析。结果表明, 每穗颖花数性状在2个组合的各分离世代均未出现超亲分离, 而其它4个性状均有不同程度的超亲分离。一次枝梗数受1对主基因+多基因控制; 其余4个性状均受2对主基因+多基因控制。每穗颖花数、每穗实粒数、穗长和二次枝梗数4个性状以主基因遗传为主, 一次枝梗数性状以多基因遗传为主。     

Statistical Analysis of Joint Effects of Major Genes and Polygenes in Quantitative Genetics

In this paper we analyze a quantitative genetic character which is controlled by both major genes and polygenes. Assuming that there are no epistatic effects, no linkage and no genetic-environmental interactions, we follow TAN and CHANG (1972) to derive the probability distributions for segregating populations. The numbers of major genes and polygenes, and the additive and dominance effects of major genes and polygenes are then estimated by using the procedures developed in TAN and CHANG (1972) and the POWELL -FLETCHER search procedure for maximum values. In this paper, we consider the case involving data from P₁, P₂, F₂, B₁ (Backcross to P₁) and B₂ (Backcross to P₂) as this type of experiment is common in practical applications. The analyses are applied to a simulated model generated by using binomial, multinomial and normal variables and to the data of an experiment on kernel weight of sorghum plant provided to the authors by Professor GEORGE H. L. LIANG of Kansas State University. The analysis of these data indicate clearly that the method derived in this paper is useful and desirable.     

Genetic Analysis of Oil Content in Brassica napus L. Using Mixed Model of Major Gene and Polygene

The joint segregation analysis of a mixed genetic model of major gene plus poly-gene was conducted to study the inheritance of oil content in Brassica napus L. Five populations, i.e the populations of 2 parents(P₁ and P₂), F₁, F₂ and F_2:3 (derived from F₂) family, from each of the two crosses (1141B × Ken C₁-1, 32B × Ken C₁-2) were investigated. The frequency distributions of oil content in F₂ and F_2:3 family populations show characteristics of a mixed normal distribution, which indicated that the inheritance of oil content followed a major gene plus poly-gene model. Twenty-one genetic models were established, which could be classified into five types: one and two major genes, polygenes, one and two major genes plus polygenes. The most suitable genetic model could be selected using Akaike's Information Criterion and the fitness of the selected one could be examined by a set of tests. Results show that genetic model D-2 is the most fitting genetic model for the trait. In other words, oil content in oilseed rape is controlled by one additive major gene plus additive and dominance polygenes. For cross 1 (1141B × Ken C₁-1) the heritabilities of major gene and poly-genes in F₂ are 68.21% and 27.17%, respectively, and in F_2:3 are 81.70% and 16.80%, respectively. The additive effect of major gene is-1.74, which indicates that the locus of the allele in parent 1141B may decrease the oil content, but that in parent Ken C₁-1 may increase it. The additive and dominance effects of the polygenes are 1.20 and -1.93, respectively. For cross 2 (32B × Ken C₁-2) the heritabilities of major gene and polygenes in F₂ are 66.20% and 28.10%, respectively, and in F_2:3 were 81.00% and 14.90%, respectively. The additive effect of major gene was -3.74, which also indicates that the locus of the allele in parent 32B may decrease the oil content, but that in parent Ken C₁-2 may increase it. The additive and dominance effects are -1.99 and 0.93, respectively. The heritability of the major gene in F_2:3 is higher than that in F₂ in both crosses, so it would be more efficent to conduct selection in F_2:3 families for high oil content in breeding.     

Genes for Dwarfness in Wheat, TRITICUM AESTIVUM L

The genetic control of plant height was studied in crosses of four spring wheats involving the standard height variety Ramona 50 and short-statured selections Olesen, D6301, and D6899. Data from parent, F₁, F₂, and F₃ populations indicated that four independently segregating loci account for most of the differences among the four varieties. Two major genes of a highly recessive nature condition reduced height in Olesen and the Norin 10 derivative D6301. Olesen also carries a third dwarfing gene which is partially dominant in its effects over genes for tallness. This gene, or a gene that acts in a similar manner, is also present in the standard height variety Ramona 50. Dwarfing in D6899, a derivative of Tom Thumb, is controlled primarily by a single gene with mainly additive effects which is not present in any of the other three varieties.

Genetic components estimated from generation means (parental, F₁, F₂, F₃, and backcross) indicated that additive gene effects were the major component of variation in four of the six crosses, and of similar magnitude to dominance effects in another cross. The primary source of genetic variation in the cross Olesen x D6899 was due to epistasis with both additive x additive and dominance x dominance effects of major importance. The results of the generation mean analyses were consistent with the models for major-gene control of plant height based on segregation patterns.

     

烟草白粉病抗性基因的遗传分析

以烟草抗白粉病品种台烟7号为母本,感病品种NC89为父本,构建6个世代的群体,利用主基因 多基因混合遗传模型的分离分析方法,研究烟草白粉病的抗性遗传规律。结果表明,烟草白粉病抗性的遗传是由两对加性-显性-上位性主基因 加性-显性-上位性多基因（E-0模型）控制的。B1、B2和F2世代主基因的遗传率分别为88.05%、32.62%、84.43%,主基因遗传率很大,说明可以在抗病育种早期进行选择;B1、F2世代多基因遗传率均为0.00%,说明烟草白粉病的发生受一定环境影响。     

Genetic analysis of waxy locus in rice (Oryza sativa L.)

Summary Inheritance of waxy locus was studied in crosses of a waxy variety with four non-waxy parents having high-, intermediate-, low- or very low-amylose content. The analysis for amylose content was done on a single grain basis in parents, F₁, F₂, B₁F₁, and B₂F₁ seeds. The waxy parent lacking synthesis of amylose content was found to differ from the ones having high-, intermediate-, low- or very low-amylose content by one gene with major effect. Dosage effects for amylose content were observed to have great influence on segregation pattern and efficiency of selection. Selection efficiency for amylose content can be enhanced by selecting for endosperm appearance in early segregating generations.     

利用P1、F1、P2、F2和F2:3家系五世代联合分离分析的拓展

在王建康等[3]的基础上拓展了利用P1、F1、P2、F2和F2：3家系5群体的2对主基因（B）和2对主基因 多基因（E）2类模型,为使拓展模型成为可能并提高分布参数估计值的精度,用IECM算法估计样本似然函数分布参数,通过重新分析3个大豆杂交组合抗豆秆黑潜蝇主茎虫量遗传资料证实了通过孟德尔氏遗传分析法所获得的结果,并得到存在多基因的统计学依据。     

Mutation loci and intragenic selection marker of the granule-bound starch synthase gene in waxy maize

Four pairs of specific PCR primers have been designed on the basis of the sequence of the granule-bound starch synthase gene (GBSS; dominant non-waxy gene Wx) and used to amplify its homologous sequence from thirteen waxy and two non-waxy inbred lines. Results from electrophoresis indicated that the recessive waxy gene was wx, derived from the dominant non-waxy gene Wx by mutation at its 3′ end. The sequence of the mutated 3′ end was amplified by the TAIL-PCR technique. Sequence alignment showed that the mutation of the wx gene was caused by transposition of the aldehyde dehydrogenase gene rf2. Two pairs of specific primers were designed on the basis of the sequence difference between the dominant gene Wx and its mutated recessive allele wx and used as intragenic selection markers to identify individual plants of genotypes WxWx, Wxwx, and wxwx by PCR amplification from the segregating population of the F₂ generation crossed between waxy and non-waxy inbred lines. Iodine solution staining and starch component assay showed that all the 35 F₂ plants identified as genotype WxWx produced non-waxy kernels of the F₃ generation and that all 33 F₂ plants identified as genotype wxwx produced waxy kernels of the F₃ generation. This result can be used to improve the selection efficiency of waxy maize breeding and for selection of other single genes and major polygenes.