植物性状-标记关联分析研究进展

要对植物改良和复杂性状进行解析,首先需要对种质多样性及其性状遗传基础进行了解.随着基因组学的发展和生物统计软件的完善,特别是分子标记技术在植物遗传育种领域的广泛应用,以连锁不平衡(LD)为基础的关联分析方法的出现为植物数量性状遗传的研究提供了新途径,也为植物的分子设计育种提供了新的思路.本文在介绍关联分析方法的基础上,综述了关联分析在植物方面的研究进展,并讨论了关联分析在植物中应用的前景.     

中国栽培和野生大豆农艺品质性状与SSR标记的关联分析 I. 群体结构及关联标记

关联作图是一种利用连锁不平衡(linkage disequilibrium, LD)检测自然群体中基因位点及其等位变异的方法。利用60个SSR标记, 对全国大豆地方品种群体(393份代表性材料)和野生大豆群体(196份代表性材料)的基因组变异进行扫描, 分析两类群体的连锁不平衡位点、群体结构, 并采用TASSEL软件的GLM (general linear model)方法对16个农艺、品质性状观测值进行标记与性状的关联分析。结果表明: (1)在公共图谱上不论共线性的或是非共线性的SSR位点组合都有一定程度的LD, 说明历史上发生过连锁群间的重组; 栽培群体的连锁不平衡成对位点数较野生群体多, 但野生群体位点间连锁不平衡程度高, 随距离的衰减慢。(2) 群体SSR数据遗传结构分析发现, 栽培群体和野生群体分别由9和4个亚群体组成, 亚群的划分与群体地理生态类型相关联, 证实地理生态类型划分有其遗传基础。(3) 栽培群体中累计有27个位点与性状相关; 野生大豆种质中累计有34个位点与性状相关。部分标记在两类群体中都表现与同一性状关联, 检出的位点有一致性, 也有互补性; 一些标记同时与2个或多个性状相关联, 可能是性状相关乃至一因多效的遗传基础; 关联位点中累计有24位点(次)与遗传群体连锁分析定位的QTL一致。     

大豆育成品种农艺性状QTL与SSR标记的关联分析

利用85个SSR标记,对大豆育成品种群体(190份代表性材料)的基因组进行扫描,在检测群体结构基础上搜索连锁不平衡位点,并采用TASSEL软件的GLM方法对11个大豆农艺性状QTL进行关联分析。结果表明：(1) 在公共图谱上共线性或非共线性的SSR位点组合均广泛存在连锁不平衡(LD),但不平衡程度D′>0.5的组合数只占总位点组合的1.71%,共线位点D′值随遗传距离衰减较快;(2) SSR数据遗传结构分析表明,育成品种群体由7个亚群体组成,矫正后全群体共有45个位点累计136个位点(次)与11个大豆农艺性状QTL关联,其中22个位点(次)与家系连锁定位的QTL区间相重,有43个位点(次) 2年重复出现;(3) 一些标记同时与2个或多个性状关联,可能是性状相关或一因多效的遗传基础;(4) 育成品种群体关联位点与地方品种群体和野生群体只有少数相同,群体间育种性状的遗传结构有相当大差异;(5) 发掘出农艺性状优异等位变异及其载体品种,包括增效最大的产量等位变异Satt347-300 (+932 kg hm^-2,中豆26),生物量等位变异Satt365-294(+3 123 kg hm^-2,黄毛豆),蛋白质含量等位变异Be475343-198 (+0.41%,淮豆4号),脂肪含量等位变异Satt150-273 (+2.32%,科丰15)等。在此基础上作了设计育种的探讨。     

全基因组关联分析在油菜遗传育种中的应用和研究进展

全基因组关联分析(genome-wide association studies,GWAS)是近几年发展起来的解析作物表型多样性遗传基础的有效分析手段。GWAS以连锁不平衡为基础鉴定某一作物群体内农艺性状即表型数据与全基因组基因型数据或候选基因间的具体关系,具有高通量(即在全基因组范围有效检测性状与基因位点的关联性)、精度高和花费时间少等显著优点,其在作物遗传育种中的作用日益凸显。本综述在系统介绍GWAS这种方法的基础上,详尽总结了其在油菜遗传育种中的应用和研究进展、存在的潜在问题及解决途径,以期为进一步利用GWAS进行油菜各种性状遗传基础的研究提供依据和参考。     

植物基因组选择及其应用研究进展

为促进基因组选择在植物遗传改良中的应用,笔者总结了基因组选择的原理与方法、分析了基因组选择育种的统计方法,指出了各种因素对基因组选择的影响,总结了基因组选择面临的挑战,并讨论了基因组选择在植物数量性状分子育种研究中可能的应用。认为随着基因型分析成本的降低和统计方法的发展,植物基因组选择将会逐步完善,将在植物基因组育种中发挥重要的作用。     

关联分析及其在果树育种研究中的应用

果树在中国农业发展中具有重要作用,而且其经济性状大多属于数量性状。基于连锁分析的QTL作图是研究这些复杂性状的有效手段,但尚存在一定的局限性。随着分子生物学的发展,关联分析已然成为发掘性状控制基因的重要手段之一,它以连锁不平衡为基础,发掘生物性状与遗传标记之间的对应关系,寻找控制性状的关键候选基因。本研究全面论述了关联分析的基本概念、原理、方法及其优越性。并综述了关联分析在果树数量性状、标记开发及功能基因挖掘方面的最新研究进展,进而,探讨关联分析在果树种质资源中的发展方向,并对其应用前景提出展望。     

基于SNP标记的关联分析在玉米耐旱研究中的应用

单核苷酸多态性(Single nucleotide polymotphisin,SNP)标记的发展促进了关联分析对植物复杂数量性状的遗传研究.研究证明:多数SNP变异与基因功能密切相关,通过关联分析可以发掘这些功能型位点变异并应用于作物遗传育种.因此,本文综述了近些年来在玉米上利用SNP标记进行关联分析的研究进展,展望了其在耐旱分子育种和遗传网络解析方面的应用前景.     

RFLP连锁图与遗传育种

RFLP标记是在分子克隆技术基础上发展起来的一种新型的遗传标记。利用这种标记可以在一个通过有性过程产生的分离群体中使用常规的遗传分析建立遗传连锁图谱,即RFLP连锁图。一旦育种上目标性状的基因被定位在RFLP图谱上,应用连锁标记的选择就能显著地提高常规育种的效率。用RFLP图谱也能定位数量性状的基因。     

大豆籽粒蛋白质含量相关QTL定位研究进展

籽粒蛋白质含量是大豆品质性状改良的主要目标之一。笔者介绍了大豆遗传图谱的构建与基因组测序发展历程,从基于分离群体的连锁分析和基于自然群体的关联分析两方面阐述了大豆籽粒蛋白质含量QTL定位研究进展,进而讨论了大豆蛋白质含量MAS育种存在的问题,最后展望了大豆蛋白质含量分子遗传改良的研究趋势。以期为大豆高蛋白育种提供参考。     

MAGIC群体的遗传特征及其在作物耐逆研究上的应用

多亲本高世代互交（multi-parent advanced generation intercross,MAGIC）群体是近年来发展起来的新一代遗传作图及育种群体。MAGIC群体最初是以研究动物及人类复杂性状遗传基础为目标而构建的基于多亲本的重组近交群体,随后将其构建方法衍生到植物中应用。MAGIC群体应用于作物遗传育种,可以建立包含无限多株系的群体,主要的优势是拥有大量可利用的多样性遗传基因池。亲本可选用育种中性状优异的材料,通过多次重组创造大量的遗传变异。群体中选出的优良株系可用作育种中间材料或直接组配新品种,也可灵活应用于数量性状位点（quantitative trait locus,QTL）的精确遗传定位分析,真正做到育种群体和定位群体的有机整合。作物耐逆性大多是由QTL控制,因此,主要围绕MAGIC群体的定义、构建流程、遗传特征及其在作物耐逆性研究上的应用和发展前景作综合性阐述。     

植物关联分析的研究进展及其在茶树分子标记辅助育种上的应用前景

茶树是我国重要的经济和饮料作物,由于自交不亲和和长期的异花授粉使其高度异质杂合,选育一个良种耗时长,提高早期选育效率和准确鉴定目标性状成为茶树育种家关心的重大问题之一。关联分析是研究分子标记或候选基因与目标性状关系的一种新方法,现已广泛应用于植物等位基因发掘、功能基因验证以及功能性标记的开发和应用等各方面的研究中,为茶树遗传育种研究提供了新思路。本文主要介绍了关联分析的基本原理、策略和及其在植物中的应用现状,并且探讨了其在茶树分子标记辅助育种研究上的应用前景。     

ICRISAT花生微核心种质农艺性状和黄曲霉抗性关联分析

以国际半干旱热带地区作物研究所(ICRISAT)花生微核心种质146份资源为品种,鉴定农艺性状和黄曲霉抗性,用26对SSR引物检测多态性位点,在分析连锁不平衡、群体结构和Kinship的基础上进行关联分析。连锁不平衡的分布显示R²平均值为0.185,表明26对SSR引物扩增的120个位点之间具有较低的连锁不平衡程度。群体结构分析结果将146份花生品种分为2个亚群,分别对应疏枝亚种和密枝亚种,与植物学分类和遗传分化分析的结果基本一致。关联分析表明,共有39个位点与10个农艺性状(株高、总分枝数、第一分枝数、小叶宽、结果分枝数、百果重、出仁率、单株生产力、种子长、种子宽)相关联,表型变异解释率为1.50%~20.34%,16个SSR位点与黄曲霉侵染病情指数、黄曲霉产毒量相关联,表型变异解释率为5.23%~17.19%,与农艺性状、黄曲霉抗性同时相关联的SSR位点有13个。关联位点的等位变异效应分析表明,10个农艺性状和2个黄曲霉抗性性状共有63个增效等位变异和47个减效等位变异,并发掘了ICG6022等携有优良等位变异的载体品种。     

Quantitative trait loci for rice blast resistance detected in a local rice breeding population by genome-wide association mapping

Plant breeding programs aim to develop cultivars with high adaptability to the specific conditions in a local region. As a result, unique genes and gene combinations have been accumulated in local elite breeding populations during the long history of plant breeding. Genetic analyses on such genes and combinations may be useful for developing new cultivars with more-desirable agronomic traits. Here, we attempted to detect quantitative trait loci (QTL) for rice blast resistance (BR) using a local breeding rice population from Hokkaido, Japan. Using genotyping data on single nucleotide polymorphisms and simple sequence repeat markers distributed throughout the whole genomic region, we detected genetic regions associated with phenotypic variation in BR by a genome-wide association mapping study (GWAS). An additional association analysis using other breeding cultivars verified the effect and inheritance of the associated region. Furthermore, the existence of a gene for BR in the associated region was confirmed by QTL mapping. The results from these studies enabled us to estimate potential of the Hokkaido rice population as a gene pool for improving BR. The results of this study could be useful for developing novel cultivars with vigorous BR in rice breeding programs.     

小麦南大2419及其衍生品种(系)主要农艺性状的关联分析

筛选与小麦重要农艺性状相关联的SSR标记,对小麦分子标记辅助育种有重要的实践意义.本研究利用多态性较高的80个SSR标记,对南大2419及其71份衍生后代品种(系)进行基因型分析,采用TASSEL软件的MLM (Mixed linear model)方法对籽粒产量、千粒重、有效穗、穗粒数等8个主要农艺性状进行SSR标记...     

Association analysis of physicochemical traits on eating quality in rice (Oryza sativa L.)

Improvement of rice eating quality is an important objective in current breeding programs. In this study, 130 rice accessions of diverse origin were genotyped using 170 SSR markers to identify marker–trait associations with physicochemical traits on eating quality. Analysis of population structure revealed four subgroups in the population. Linkage disequilibrium (LD) patterns and distributions are of fundamental importance for genome-wide mapping associations. The mean r ² value for all intrachromosomal loci pairs was 0.0940. LD between linked markers decreased with distance. Marker–trait associations were investigated using the unified mixed-model approach, considering both population structure (Q) and kinship (K). In total, 101 marker–trait associations (p < 0.05) were identified using 52 different SSR markers covering 12 chromosomes. The results suggest that association mapping in rice is a viable alternative to quantitative trait loci mapping, and detection of new marker–trait associations associated with rice eating quality will also provide important information for marker-assisted breeding and functional analysis of rice grain quality.     

Genome-wide association mapping focusing on a rice population derived from rice breeding programs in a region

Plant breeding programs in local regions may generate genetic variations that are desirable to local populations and shape adaptability during the establishment of local populations. To elucidate genetic bases for this process, we proposed a new approach for identifying the genetic bases for the traits improved during rice breeding programs; association mapping focusing on a local population. In the present study, we performed association mapping focusing on a local rice population, consisting of 63 varieties, in Hokkaido, the northernmost region of Japan and one of the northern limits of rice cultivation worldwide. Six and seventeen QTLs were identified for heading date and low temperature germinability, respectively. Of these, 13 were novel QTLs in this population and 10 corresponded to the QTLs previously reported based on QTL mapping. The identification of QTLs for traits in local populations including elite varieties may lead to a better understanding of the genetic bases of elite traits. This is of direct relevance for plant breeding programs in local regions.     

Quantitative trait loci for agronomic traits in soybean

There continues to be improvement in seed yields of soybean by conventional breeding, but molecular techniques may provide faster genetic gains. The objective of this study was to identify quantitative trait loci (QTL) associated with the agronomic traits seed yield, lodging, plant height, seed filling period and plant maturity in soybean. To achieve this objective, 101 F₆‐derived recombinant inbred lines (RIL) from a population developed from a cross of N87‐984‐16 × TN93‐99 were used. Experiments were conducted in six environments during 2002–2003. Heritability estimates on an entry mean basis from data combined across environments ranged from 0.12 to 0.65 for seed yield and seed filling period, respectively. Composite interval mapping detected one QTL for yield (near Satt076), two for lodging (near Satt225 and Satt593) and four for maturity (near Satt263, Satt292, Satt293 and Satt591) in this population. Additional environmentally sensitive QTL for these traits, and for seed filling period and plant height are also reported. The QTL associated with agronomic traits that we report and the recently released germplasm (PI 636460) from this population may be useful in soybean breeding programmes.     

甘蓝型油菜产量及相关性状的QTL分析

高产是甘蓝型油菜育种的重要目标之一，产量是多基因控制的数量性状。本文通过QTL作图分析了产量及其相关性状的数量性状位点，以甘蓝型油菜中油821和保604 F₁代小孢子培养获得的DH系为作图群体，构建了由20个连锁群组成的，包括251个分子标记( 2个RFLP标记，72个RAPD标记，91个SSR标记，86个SRAP标记)的遗传连锁图(10个标记没有分配到连锁群中)。图谱的平均图距为6.96 cM，共覆盖油菜基因组1 746.5 cM。在此图谱基础上采取复合区间作图法，检测到与油菜产量及其相关性状有关的QTL共17个。其中控制株高的3个分别位于第4、第9和第10连锁群上，对性状的解释率为9.42%～17.58%；与分枝部位有关的4个分别位于第4、第6和第7连锁群上，其中Bp1 和Bp2 均位于第4连锁群，对性状的解释率为8.13%～15.20%；与主花序有效长有关的3个分别位于第4、第10和第16连锁群上，对性状的解释率为7.49%～23.36%；与一次有效分枝有关的2个分别位于第1、第4连锁群上，对性状的解释率为15.29%～19.58%；与角果总数和千粒重有关的分别位于第4连锁群和第9连锁群上，贡献率分别为17.42%和7.64%；与单株产量有关的3个分别位于第3、第4和第15连锁群，共解释26.60%的表型变异。部分性状的QTL在连锁群上成簇分布,对性状贡献率很大，表现主效QTLs的特点，相应的性状之间也呈显著相关，这表明一因多效或者相关的QTLs之间紧密连锁是性状相关的遗传基础。本研究中与主效QTLs连锁的标记可用于油菜产量性状的分子标记辅助选择。     

Association Analysis of Agronomic and Yield-Related Traits in an Upland Cotton (Gossypium hirsutum L.) Germplasm Population Based on SSR Markers

[Objective] Molecular markers associated with agronomic and yield-related traits could provide useful references for improving the efficiency of Gossypium hirsutum breeding. [Method] In 147 upland cotton materials, 237 Simple Sequence Repeat(SSR) markers were assayed for polymorphisms. The population structure was analyzed using Structure 2.2. An association analysis between SSR markers and the agronomic and yield-related traits was performed using the Tassel 2.1 General Linear Model program. [Result] The average coefficients of variation for the five indicators varied from 7.28% to 21.06%, with an average of 13.47%, which indicated that the test population had a rich phenotypic diversity. The 237 SSR markers included 690 alleles belonging to 281 loci, with an average of 2.455 0 alleles per SSR marker (ranging from 2 to 6), and the average polymorphism information content value was 0.216 5. The test group could be divided into seven subgroups using Structure software. A genome-wide association analysis was conducted on five agronomic and yield-related traits from 2013 to 2015. There were 45 SSR loci and 1 locus significantly (P＜0.01) associated with boll weight and lint percentage, which were detected in 3 or more environments, and they explained 19.30% and 11.58%, respectively, of the phenotypic variation in the different environments (P＜0.01). Nine loci were significantly (P＜0.01) associated with agronomic traits at two or more bowing times, and one, four and four markers were associated with fruit branch number, boll number and plant height, explaining 9.96%, 7.00% and 5.57%, respectively, of the phenotypic variation in different environments. Sites significantly associated with boll weight, fruit branch number and plant height were simultaneously detected near BNL2448. [Conclusion] Many loci associated with the yield and other important agronomic traits of upland cotton were repeatedly detected in different environments using an association analysis. The results provide useful reference information for cotton molecular marker-assisted selection-based breeding.