Improving breeding efficiency in potato using molecular and quantitative genetics

Key message

Potatoes are highly heterozygous and the conventional breeding of superior germplasm is challenging, but use of a combination of MAS and EBVs can accelerate genetic gain.

Abstract

Cultivated potatoes are highly heterozygous due to their outbreeding nature, and suffer acute inbreeding depression. Modern potato cultivars also exhibit tetrasomic inheritance. Due to this genetic heterogeneity, the large number of target traits and the specific requirements of commercial cultivars, potato breeding is challenging. A conventional breeding strategy applies phenotypic recurrent selection over a number of generations, a process which can take over 10 years. Recently, major advances in genetics and molecular biology have provided breeders with molecular tools to accelerate gains for some traits. Marker-assisted selection (MAS) can be effectively used for the identification of major genes and quantitative trait loci that exhibit large effects. There are also a number of complex traits of interest, such as yield, that are influenced by a large number of genes of individual small effect where MAS will be difficult to deploy. Progeny testing and the use of pedigree in the analysis can provide effective identification of the superior genetic factors that underpin these complex traits. Recently, it has been shown that estimated breeding values (EBVs) can be developed for complex potato traits. Using a combination of MAS and EBVs for simple and complex traits can lead to a significant reduction in the length of the breeding cycle for the identification of superior germplasm.     

作物分子标记辅助选择的研究进展、影响因素及其发展策略

随着分子标记技术及其检测手段的发展,开发和应用成本的降低,分子标记辅助选择(MAS)在作物育种上的应用优势日益明显。本文综述了近年来MAS在基因聚合、基因转移和数量性状改良上的研究进展。总结了MAS的影响因素,包括标记与基因间的距离、目标性状的遗传率、群体大小、所用分子标记的数目、类型和相位等。并提出育种和定位同步进行、选择合适分子标记类型和数量、简化DNA提取方法、背景选择的逐步选择法、确定合适选择方案等MAS发展策略。     

Efficiency of Marker-Assisted Selection in the Improvement of Quantitative Traits

Molecular genetics can be integrated with traditional methods of artificial selection on phenotypes by applying marker-assisted selection (MAS). We derive selection indices that maximize the rate of improvement in quantitative characters under different schemes of MAS combining information on molecular genetic polymorphisms (marker loci) with data on phenotypic variation among individuals (and their relatives). We also analyze statistical limitations on the efficiency of MAS, including the detectability of associations between marker loci and quantitative trait loci, and sampling errors in estimating the weighting coefficients in the selection index. The efficiency of artificial selection can be increased substantially using MAS following hybridization of selected lines. This requires initially scoring genotypes at a few hundred molecular marker loci, as well as phenotypic traits, on a few hundred to a few thousand individuals; the number of marker loci scored can be greatly reduced in later generations. The increase in selection efficiency from the use of marker loci, and the sample sizes necessary to achieve them, depend on the genetic parameters and the selection scheme.     

Germplasm-regression-combined (GRC) marker-trait association identification in plant breeding: a challenge for plant biotechnological breeding under soil water deficit conditions

In the past 20 years, the major effort in plant breeding has changed from quantitative to molecular genetics with emphasis on quantitative trait loci (QTL) identification and marker assisted selection (MAS). However, results have been modest. This has been due to several factors including absence of tight linkage QTL, non-availability of mapping populations, and substantial time needed to develop such populations. To overcome these limitations, and as an alternative to planned populations, molecular marker–trait associations have been identified by the combination between germplasm and the regression technique. In the present preview, the authors (1) survey the successful applications of germplasm–regression–combined (GRC) molecular marker–trait association identification in plants; (2) describe how to do the GRC analysis and its differences from mapping QTL based on a linkage map reconstructed from the planned populations; (3) consider the factors that affect the GRC association identification, including selections of optimal germplasm and molecular markers and testing of identification efficiency of markers associated with traits; and (4) finally discuss the future prospects of GRC marker–trait association analysis used in plant MAS/QTL breeding programs, especially in long-juvenile woody plants when no other genetic information such as linkage maps and QTL are available.     

Whole-genome strategies for marker-assisted plant breeding

Molecular breeding for complex traits in crop plants requires understanding and manipulation of many factors influencing plant growth, development and responses to an array of biotic and abiotic stresses. Molecular marker-assisted breeding procedures can be facilitated and revolutionized through whole-genome strategies, which utilize full genome sequencing and genome-wide molecular markers to effectively address various genomic and environmental factors through a representative or complete set of genetic resources and breeding materials. These strategies are now increasingly based on understanding of specific genomic regions, genes/alleles, haplotypes, linkage disequilibrium (LD) block(s), gene networks and their contribution to specific phenotypes. Large-scale and high-density genotyping and genome-wide selection are two important components of these strategies. As components of whole-genome strategies, molecular breeding platforms and methodologies should be backed up by high throughput and precision phenotyping and e-typing (environmental assay) with strong support systems such as breeding informatics and decision support tools. Some basic strategies are discussed in this article, including (1) seed DNA-based genotyping for simplifying marker-assisted selection (MAS), reducing breeding cost and increasing scale and efficiency, (2) selective genotyping and phenotyping, combined with pooled DNA analysis, for capturing the most important contributing factors, (3) flexible genotyping systems, such as genotyping by sequencing and arraying, refined for different selection methods including MAS, marker-assisted recurrent selection and genomic selection (GS), (4) marker-trait association analysis using joint linkage and LD mapping, and (5) sequence-based strategies for marker development, allele mining, gene discovery and molecular breeding.     

动物遗传标记辅助选择研究及其应用

随着分子数量遗传学及其相关学科的发展,有关动物遗传标记辅助选择方面的研究也在不断深入,且已经在动物遗传改良中有了一些成功应用的示例。就如何综合利用表型、系谱和遗传标记信息进行育种值估计的统计学方法研究方面,目前已基本形成了较为完善的统计学方法。同时,在标记辅助选择相对效率及其影响因素,以及标记辅助选择实施方案的研究上也取得了不少喜人的成果。本文综述了动物遗传标记辅助选择研究的一些进展,并对标记辅助选择在动物遗传改良中应用的有关问题进行了讨论。 Abstract:With the development of molecular and quantitative genetics and its related subjects,it made a great progress on the research about animal genetic marker-assisted selection (MAS).There were also some successful examples on the application of MAS to animal genetic improvement.The statistical method which using phenotypic,pedigree and genetic marker information to predict individual breeding values has already been developed.Many achievements were obtained from the researches,which carried on MAS relative efficiency and its affecting factors and selection schemes.The present paper reviewed some progresses of MAS research and discussed some problems about MAS application to animal breeding.     

Usefulness of marker‐assisted selection to improve maize for increased resistance to Sesamia nonagrioides attack with no detrimental effect on yield

Two decades of investigations on maize resistance to Mediterranean corn borer (Sesamia nonagrioides Lefebvre; MCB) have shown that breeding for increased resistance to stem tunnelling by MCB often resulted in reduced yield because significant genetic correlation between both traits exists in some backgrounds. Unlike phenotypic selection, marker‐assisted selection (MAS) could differentiate markers linked only to one trait from those linked simultaneously to yield potential and susceptibility to the pest. In the current study, the suitability of MAS for improving resistance to stem tunnelling without adverse effects on yield has been tested. The unfavourable genetic relationship between yield potential and susceptibility could be overcome using MAS. Gains obtained using MAS were weak, because genetic variance explained by the quantitative trait loci (QTL) was low but results encourage us to persevere in using marker information for simultaneous improvement of resistance and yield especially if genome‐wide approaches are applied. Approaches to detect QTL are widely used, but studies on the suitability of markers linked to QTL for performing MAS have been mostly neglected.     

Marker-assisted selection with spatial analysis of unreplicated field trials

 Many studies have shown that molecular markers can improve the efficiency of the selection of quantitative traits in plant breeding provided that large population sizes are used. As a way to limit experimental costs it appears that the use of unreplicated trials may be more valuable than the use of replicated plots in one trial. In this particular context of unreplicated large trials, spatial heterogeneity within the field may reduce the efficiency of the selection. The problem of controlling spatial heterogeneity was seldom considered in the case of marker-assisted selection (MAS). Here, we propose an integrated method to predict genetic values considering simultaneously marker information and possible spatial heterogeneity. This method was applied to a population of 300 F₃ lines of maize evaluated in 11 unreplicated trials for grain yield. We show that when spatial field heterogeneity is considered through appropriate statistical models the accuracy of genetic value predictions is improved and the same genetic gain can be achieved with a reduced number of trials. Received: 7 January 1998 / Accepted: 20 July 1998     

Genetic and genomic analyses for economically important traits and their applications in molecular breeding of cultured fish

The traits of cultured fish must continually be genetically improved to supply high-quality animal protein for human consumption.Economically important fish traits are controlled by multiple gene quantitative trait loci(QTL),most of which have minor effects,but a few genes may have major effects useful for molecular breeding.In this review,we chose relevant studies on some of the most intensively cultured fish and concisely summarize progress on identifying and verifying QTLs for such traits as growth,disease and stress resistance and sex in recent decades.The potential applications of these major-effect genes and their associated markers in marker-assisted selection and molecular breeding,as well as future research directions are also discussed.These genetic and genomic analyses will be valuable for elucidating the mechanisms modulating economically important traits and to establish more effective molecular breeding techniques in fish.     

Prospects for increasing yield in macadamia using component traits and genomics

Selection of candidate cultivars in macadamia requires extensive phenotypic measurements over many years and trials. In particular, yield traits such as nut-in-shell yield and kernel yield are economically vital characteristics and therefore guide the selection process for new cultivars. However, these traits can only be measured in mature trees, resulting in long generation intervals and slow rates of genetic gain. In addition, these traits are expensive to measure. Strategies to reduce the generation interval and increase the intensity of selection include using yield component traits, identification of markers associated with component traits, and genomic selection for yield. Yield component traits that contribute to resource availability for fruit formation include floral and nut characteristics. In this review, these traits will be investigated to estimate their relative importance in macadamia breeding and their heritability and correlations with yield. Furthermore, the usefulness of genome-wide association studies regarding yield component traits will be reviewed. Genetic-based breeding techniques could exploit this information to increase yield gains per breeding cycle and estimate the quantitative nature of yield traits. Genomic selection uses genome-wide molecular markers to predict the phenotype of individuals at an early age before maturity, thereby reducing the cycle time and increasing gain per unit time in plant breeding programmes. This review evaluates the potential for measurement of yield component traits, genome-wide association studies, and genomic selection to be employed in the Australian macadamia breeding programme to accelerate gains for nut yield.     

H-FABP、MC4R、ADD1基因多态性在3个猪群中分布及其与肌内脂肪和背膘的相关研究

具有不同遗传特性的猪种具有不同的肉质性状。尤其是地方品种和引进品种间在肉质性状存在极大的差异。在已有的研究中H-FABP,MC4R,ADD1基因同肌内脂肪或背膘相关。利用梅山猪、苏太猪和杜×长×大猪为试验动物,研究上述3个基因的多态性分布和多态性同肌内脂肪和背膘的相关性。结果表明：3个基因的多态性分布在不同猪种间存在极显著的差异,这种差异可能是肌内脂肪（IMF）或背膘（BF）不同的主要原因之一。连锁分析表明：H-FABP和ADD1基因多态性同IMF有显著的相关,但是同BF没有显著的相关;MC4R基因的多态性同IMF和BF都有显著相关性。说明：H-FABP和ADD1基因多态性有可能应用到提高IMF,同时不影响BF的育种实践中。      

性状遗传力与QTL方差对标记辅助选择效果的影响

在采用动物模型标记辅助最佳线性无偏预测方法对个体育种值进行估计的基础上,模拟了在一个闭锁群体内连续对单个性状选择10个世代的情形,并系统地比较了性状遗传力和QTL方差对标记辅助选择所获得的遗传进展、QTL增效基因频率和群体近交系数变化的影响。结果表明：在对高遗传力和QTL方差较小的性状实施标记辅助选择时,可望获得更大的遗传进展;遗传力越高,QTL方差越大,则QTL增效基因频率的上升速度越快;遗传力较高时,群体近交系数上升的速度较为缓慢,而QTL方差对群体近交系数上升速度的影响则不甚明显。结合前人关于标记辅助选择相对效率的研究结果,可以认为：当选择性状的遗传力和QTL方差为中等水平时,标记辅助选择可望获得理想的效果。     

标记辅助回交育种中所需最小样本容量的近似估计

回交育种是把有利基因从供体亲本向受体亲本转移的一种有效方法,标记辅助选择可加速其进程。为了制定合理的标记辅助选择计划,育种家必须知道所需的后代群体大小。该文提出了一种估算在标记辅助回交育种中同时进行前景选择和背景选择所需群体大小的方法。在假定所需转移的目标基因座与遗传背景之间为相互独立的简化假设下,可以通过将解析方法（针对前景选择）与基于回交亲本图示基因型的模拟方法（针对背景选择）相结合,近似地估计出在每一世代中选到所需基因型的概率,进而估算出在一定概率水平下至少获得一个符合要求的个体所需的最小样本容量,用假想的例子演示了该方法的使用情况。该方法可以很方便地应用于实际的回交育种。     

Genome-wide association and genomic selection in animal breeding

Results from genome-wide association studies in livestock, and humans, has lead to the conclusion that the effect of individual quantitative trait loci (QTL) on complex traits, such as yield, are likely to be small; therefore, a large number of QTL are necessary to explain genetic variation in these traits. Given this genetic architecture, gains from marker-assisted selection (MAS) programs using only a small number of DNA markers to trace a limited number of QTL is likely to be small. This has lead to the development of alternative technology for using the available dense single nucleotide polymorphism (SNP) information, called genomic selection. Genomic selection uses a genome-wide panel of dense markers so that all QTL are likely to be in linkage disequilibrium with at least one SNP. The genomic breeding values are predicted to be the sum of the effect of these SNPs across the entire genome. In dairy cattle breeding, the accuracy of genomic estimated breeding values (GEBV) that can be achieved and the fact that these are available early in life have lead to rapid adoption of the technology. Here, we discuss the design of experiments necessary to achieve accurate prediction of GEBV in future generations in terms of the number of markers necessary and the size of the reference population where marker effects are estimated. We also present a simple method for implementing genomic selection using a genomic relationship matrix. Future challenges discussed include using whole genome sequence data to improve the accuracy of genomic selection and management of inbreeding through genomic relationships.     

70个水稻微卫星标记染色体位置的更正

微卫星标记(SSR)因其操作简单和稳定可靠的特点而成为一种重要的分子标记,被广泛应用于遗传作图和种质鉴定等方面。但其在染色体上位置的正确性将直接影响到基因定位的正确性和后续研究的方向。利用美国国家生物信息技术中心(NCBI)网站的Blast程序,将2740个SSR标记的前后引物序列与水稻粳稻品种日本晴基因组进行比对,共发现70个标记位于另一条染色体,对这70个标记重新锚定的染色体进行了更正。这将有助于今后水稻分子标记遗传连锁图的正确构建。     

黄瓜分子标记辅助育种研究进展

本文综述了不同分子标记技术在黄瓜遗传连锁图谱构建、重要性状相关基因的定位、种质资源遗传多样性分析和亲缘关系鉴定、分子标记辅助选择、种子纯度与活力鉴定及其在黄瓜遗传育种等方面的应用,讨论了目前黄瓜遗传育种中应用分子标记技术存在的问题和今后育种工作的重点,并对黄瓜分子标记辅助育种的前景作了展望。     

Within-family marker-assisted selection for aquaculture species

A within-family marker-assisted selection scheme was designed for typical aquaculture breeding schemes, where most traits are recorded on sibs of the candidates. Here, sibs of candidates were tested for the trait and genotyped to establish genetic marker effects on the trait. BLUP breeding values were calculated, including information of the markers (MAS) or not (NONMAS). These breeding values were identical for all family members in the NONMAS schemes, but differed between family members in the MAS schemes, making within-family selection possible. MAS had up to twice the total genetic gain of the corresponding NONMAS scheme. MAS was somewhat less effective when heritability increased from 0.06 to 0.12 or when the frequency of the positive allele was < 0.5. The relative efficiency of MAS was higher for schemes with more candidates, because of larger fullsib family sizes. MAS was also more efficient when male:female mating ratio changed from 1:1 to 1:5 or when the QTL explained more of the total genetic variation. Four instead of two markers linked to the QTL increased genetic gain somewhat. There was no significant difference in polygenic genetic gain between MAS and NONMAS for most schemes. The rates of inbreeding were lower for MAS than NON-MAS schemes, because fewer full-sibs were selected by MAS.     

荔枝与龙眼种质资源研究进展

荔枝和龙眼是中国著名的南亚热带果树。本文分别就荔枝、龙眼种质资源在起源与分布、分类与鉴定等方面取得的研究进展进行了综述。荔枝和龙眼的分类研究大体经历了3个阶段:根据形态特征进行分类、应用同工酶分析和DNA分子标记进行分类。形态特征常常受到环境条件的影响,而同工酶分析可以检测到的多态性位点十分有限。应用分子标记研究荔枝、龙眼种质资源的遗传多样性克服了前述2种研究方法的不足。目前,荔枝和龙眼的品种选育仍然以实生选种和芽变选种为主。为了实现对荔枝、龙眼果实品质和农艺性状的改良,建议在分子水平上进一步深入地探讨荔枝、龙眼种质资源的遗传多样性;同时通过应用分子标记辅助选择技术提高荔枝和龙眼杂交育种的效率。     

Marker-assisted wheat breeding: present status and future possibilities

Wheat production and productivity in the past witnessed a remarkable growth. However, this growth rate could not be sustained during the last decade, causing concern among world wheat community. Marker-assisted selection (MAS), which is being practiced for improvement of a variety of traits in wheat around the world, may at least partly help in providing the desired solution. Marker-trait associations are now known for a number of simple, but difficult-to-score traits, so that MAS has been found useful for improvement of several of these important economic traits. Breeding strategies including marker-assisted backcrossing, forward breeding, MAS involving doubled haploid technology and F₂ enrichment have been successfully utilized for this purpose. However, for improvement of complex polygenic traits, newer technologies based on high throughput genotyping associated with new marker systems (e.g., DArT and SNP), and new selection strategies such as AB-QTL, mapping-as-you-go, marker-assisted recurrent selection and genome-wide selection will have to be tried in future. The progress made in all these aspects of marker-assisted wheat breeding, and the limitations and future prospects of this emerging technology have been reviewed in this article.     

不同QTL增效基因初始频率下标记辅助选择的效果

采用随机模拟方法模拟了在一个闭锁群体内连续对单个性状选择10个世代的情形。在假定选择性状受一个位于常染色体上的QTL和多基因共同控制的情况下,采用动物模型标记辅助最佳线性无偏预测方法估计个体育种值并据此进行种畜的选留,并在此基础上系统地比较了QTL增效基因初始频率对标记辅助选择效果的影响。结果表明：当群体中QTL增效基因的初始频率较低时,选择所获得的QTL基因型值的进展会更大,标记辅助选择在单位时间内可获得较大的遗传进展;此时,尽管QTL增效基因在群体中固定所需的世代数会更长一些,但其频率上升的速度却更快。而QTL增效基因初始频率的高低对群体近交增量的影响不是很大。