作物数量性状(QTL)基因研究进展

从作物数量性状基因座QTL(quantitative trait locus)作图群体类型及特点,QTL定位的原理和方法,作物QTL研究现状,以及QTL精细定位、克隆、利用等方面进行了综述。对作物QTL分子标记辅助选择育种进行了探讨,并对目前QTL定位中存在的问题和今后QTL的研究方向提出了一些思考。     

基于基因型选择提高QTL作图的精度——以一个RIL群体为例

以PCR为基础的分子标记以及其他检测技术的发展,使得大规模的标记分析成为现实。这也为通过大群体标记分析,然后基于基因型选择挑选合适的小群体,从而提高QTL定位准确性和精度提供了可能。以一个包含294个家系的重组自交系（RIL）群体为例,通过基因型选择和随机选择的办法产生了一系列大小不等的亚群体,比较了两类群体QTL定位的结果。分析表明：相同大小的基因型选择群体与随机群体相比性状的表型分布都符合正态分布;标记的偏分离情况也没有明显的差别,都随着群体大小的增大,偏分离的比例也逐渐增大。但同等大小的基因型选择群体比随机群体的交换富集率（CE）要大,且随着选择强度的增大不断增大,如群体大小为270时,CE=1．04,群体大小为30时,CE=1．45。总体上,随着群体大小的增加,不管是随机群体还是选择群体,其QTL检测能力、灵敏性和特异性也随之增加,但选择群体的检测能力、灵敏性和特异性总体上要好于随机群体。当群体大于或等于240时,其QTL检测能力基本没有差别;群体大小大于或等于210时,其QTL检测的灵敏性和特异性也没有什么差别。这也说明：选择强度越大,效果越明显。以QTLI—LOD区间作为衡量QTL精度的一个指标,结果显示所有基因型选择群体都比相同大小随机群体的QTL定位精度高。目前QTL定位研究中,基因型数据较表型数据而言更容易准确获得,因此通过基因型选择可以更好的优化群体结构,减少田间实验的工作量,提高全基因组水平QTL作图的精度,为随后的QTL辅助选择和精细定位以及克隆提供帮助。     

小麦株高性状的QTL分析

自20世纪60年代农林10号矮秆基因被用于小麦育种以来,矮化育种成为世界范围内势不可挡的趋势,矮秆基因研究被越来越多的育种专家重视,先后鉴定出20余个矮秆基因,并应用其中6,7个基因,培育了大批丰产潜力大的半矮秆品种,应用矮秆冬小麦吕系DN3338（♀）和F390（♂）杂交得到的F2：3群体,研究小麦株高的遗传基础,以控制株高的数量性状基因座进行定位,利用240个F2：3家系,构建了含215个微卫星标记,覆盖3600cM,由21个连锁群组成的遗传 连锁图谱,并对该群体进行了4个环境（2年：2000年和2001年,2点：北京和石家庄）3重复的田间种植;采用区间作图法,对该群体的株高性状进行了QTL分析。结果表明：7个影响株高的QTL分别位于染色体1B,4B(2个）,6A（2个）,6D和7A上,每个QTL能解释5．2％－50．1％的表型变异,每个环境条件下检测出的所有QTL能解释64．8％－75％,的表型变异,除了7A上的QTL外,其他6个降低株高的QTL均来自ND3338,其效应介于0．94cm-9.33cm之间,且其中的4个在所有的环境下都能被检测出来,具有较高的稳定性,在4BS的Xgwm113标记附近有一主效QTL,其在不同的环境下能降低株高7．91cm-9.33cm,解释27．8％－36．2％,的表型变异,有着同农林10号中Rht-Blb相近的效应;同时在4BS上还发现一个和地点互作的QTL,该QTL在石家庄的两年试验中均被检测到,且有较大的效应值（80cm和7.6cm),因此,认为大部分的QTL能在所有的环境中检测到,这些QTO可以被用于品种改良和分子标记辅助选择育种。     

水稻QTL分析的研究进展

水稻许多重要的性状是由多基因控制的数量性状,经典的数量遗传学只能把数量性状作为一个整体进行研究。近年来．高密度分子标记连锁图的构建和有效的生物统计学方法的发展使人们对数量性状遗传基础的研究出现了革命性的变化。通过对不同群体内的个体或品系的分子标记基因型和表型数据的共分离分析,能对QTL进行检测和定位。本文对QTL定位的原理和方法进行了介绍,从QTL的数目和效应、上位性效应、QTL基因型与环境的互作、相关性状的QTL以及个体发育不同阶段的QTL等方面对水稻QTL分析的研究进展进行了综述。水稻基因组测序计划已经完成,本文还对基因组时代水稻QTL精细定位和克隆的方法进行了探讨,对QTL分析在水稻育种中的应用前景进行了展望。     

西瓜果实性状QTL定位及其遗传效应分析

用可溶性固形物含量高、薄皮、感枯萎病的栽培西瓜自交系（Ｃｉｔｒｕｌｌｕｓ ｌａｎａｔｕｓ ｖａｒ．ｌａｎａｔｕｓ）９７１０３和可溶性固形物含量低、皮厚、抗病的野生西瓜种质（Ｃｉｔｒｕｌｌｕｓ ｌａｎａｔｕｓ ｖａｒ．ｃｉｔｒｏｉｄｅｓ）ＰＩ２９６３４１杂交所得Ｆ２的１１８个单株为作图群体,通过构建分子连锁图谱,对西瓜主要果实性状可溶性固形物含量、果皮硬度、果皮厚度、单果重、种子千粒重进行区间作图,     

BC群体不同连锁模式分子区间标记QTL作图相关方法的精确度分析

该文分析了BC群体不同连锁模式分子区间标记QTL作图相关方法的精确度,提出了相应参数的适用范匿,连锁顺序的检测方法,分析步骤,为QTL作图提供了理论基础。     

雄性不交换条件下F2群体高密度分子区间标记QTL的精确作图

提出雄性不交换条件下F2群体间标记定位QTL的相关方法,研究高密度分子标记存在强烈交叉干涉时,QTL的精确定位方法。     

豌豆遗传图谱构建及QTL定位研究进展

豌豆的许多性状是多基因控制的数量性状,QTL定位就是以分子标记技术为工具、以遗传连锁图谱为基础、利用分子标记与QTL之间的连锁关系确定控制数量性状的基因在基因组中的位置.本文对QTL定位原理、方法进行了简单介绍;对豌豆遗传图谱构建及主要性状,如产量、品质、抗病性等QTL定位、遗传效应分析等方面的研究进行综述;对目前基于QTL豌豆分子标记育种存在的问题、应用前景进行了探讨.     

分子标记辅助聚合两个棉纤维高强主效QTLs的选择效果

利用长江流域推广品种泗棉3号和优异纤维种质系7235为育种亲本,配置了系统育种和修饰回交聚合育种两套群体。基于来自7235的2个高强纤维主效QTL的分子标记,在上述育种群体中进行了分子标记辅助选择效率研究。高强纤维主效QTLfs1是利用(7235×TM1)F2分离群体,通过集团混合分离法检测到的,它可解释纤维强度表型变异的30%以上。高强纤维主效QTLfs2最初是利用(HS42710×TM1)F2分离群体检测到的,它可解释纤维强度表型变异的12.5%以上。进一步的研究表明,该QTL也位于7235优质系中,但与QTLfs1非等位。2套育种分离群体的2个高强纤维主效QTL的分子标记辅助选择效果表明:QTLfs1在不同环境条件下均稳定表达,它对不同遗传背景的育种群体均有显著的选择效果。尽管QTLfs2的选择效果低于QTLfs1,它在高世代育种群体中也表现较高的选择效率。利用分子标记辅助选择具有一定遗传距离的QTLfs1区间,其纤维强度的选择效率将大大增强。通过分子标记对位于不同连锁群上的2个QTL聚合选择,其中选单株的纤维强度显著提高。研究结果为利用分子标记辅助聚合优质QTL提供了成功实例。     

利用相关性分析鉴定与水稻根部性状表达相关的分子标记

84个水稻品种在营养液中生长,10天后测定每一品种的最大根长（Maximum Root Length,MRL)和根干重（Root Dry Weight,RDW)。选取其中有代表性的27个水稻品种,用扩增片段长度多态性（Amplified Fragment Length Polymor-phism,AFLP)技术进行基因组差异分析,通过计算差异带与性状表现间的相关系数,筛选与苗期水稻最大根长和根干重显著相关的分子标记,经过15对AFLP引物的筛选,有4对引物的7个片段的基因型表现与最大根长或（和）根干重显著相关,对其中的片段之一“T3P3f”进行克隆,测序后,设计特异PCR扩增引物“Z336”,进一步对84个水稻品种进行鉴定,统计分析后发现,Z2336与最大根长的相关系数为-0.193,相关性几近显著水平;与根干重的相关系数为-0.391,相关性达极显著水平,计算对根干重的差异解释率,可达15.3%,显示该标记与控制根干重性状表达的某个数量基因紧密连锁,它的存在对性状值的降低有显著的关系,进一步利用源于ZYQ8和JX17的加倍单倍体（double haploid,DH)分离群体进行基因定位,发现Z336位于水稻第11号染色体上,距离相邻的分子标记9.4cM。     

加性基因效应与分子标记回归方程的关系

控制数量性状的基因作用历来是遗传学工作者所关注的重要课题．本文对以正交表形式表现的共显性动物分子标记资料,根据加性效应基因控制的数量性状遗传模型配合了动物分子标记回归方程通式．结果表明：对以正交表形式表现的加性效应基因共显性分子标记资料配合的分子标记回归方程,可对加性等位基因的相对作用差加以估计,并可作为育种的依据．     

Advanced backcross QTL analysis in a cross between an elite processing line of tomato and its wild relative L. pimpinellifolium

Approximately 170 BC2 plants from a cross between an elite processing inbred (recurrent parent) and the wild species Lycopersicon pimpinellifolium LA1589 (donor parent) were analyzed with segregating molecular markers covering the entire tomato genome. Marker data were used to identify QTLs controlling a battery of horticultural traits measured on BC2F1 and BC3 families derived from the BC2 individuals. Despite its overall inferior appearance, L. pimpinellifolium was shown to possess QTL alleles capable of enhancing most traits important in processing tomato production. QTL-NIL lines, containing specific QTLs modifying fruit size and shape, were subsequently constructed and shown to display the transgressive phenotypes predicted from the original BC2 QTL analysis. The potential of exploiting unadapted and wild germplasm via advanced backcross QTL analysis for the enhancement of elite crop varieties is discussed.     

小麦数量性状分子标记的研究进展

小麦的许多重要经济性状是受多基因控制的数量性状。综述了近年来小麦QTL的研究进展,包括QTL定位原理、研究涉及的性状、QTL分布情况、贡献率、数据统计分析使用的方法和应用软件等,列举了小麦重要农艺性状QTL在育种中的应用实例,分析了小麦QTL分子标记的发展和应用前景。     

QTL analysis of an advanced backcross of Lycopersicon peruvianum to the cultivated tomato and comparisons with QTLs found in other wild species

 A BC₃ population previously developed from a backcross of Lycopersicon peruvianum, a wild relative of tomato, into the cultivated variety L. esculentum was analyzed for QTLs. Approximately 200 BC₄ families were scored for 35 traits in four locations worldwide. One hundred and sixty-six QTLs were detected for 29 of those traits. For more than half of those 29 traits at least 1 QTL was detected for which the presence of the wild allele was associated with an agronomically beneficial effect despite the inferior phenotype of the wild parent. Eight QTLs for fruit weight could be followed through the BC₂, BC₃, and BC₄, generations, supporting the authenticity of these QTLs. Comparisons were made between the QTLs found in this study and those found in studies involving two other wild species; the results showed that while some of these QTLs can be presumed to be allelic, most of the QTLs detected in this study are ones not previously discovered. Received: 9 April 1997 / Accepted: 20 May 1997     

Moving from QTL experimental results to the utilization of QTL in breeding programmes

Results from quantitative trait loci studies cannot be readily implemented into breeding schemes through marker assisted selection because of uncertainty about whether the quantitative trait loci identified are real and whether the identified quantitative trait loci are segregating in the breeding population. The present paper outlines and discusses strategies to reduce uncertainty in the results from quantitative trait loci studies. One strategy to confirm results from quantitative trait loci studies is to combine P -values from many quantitative trait loci experiments, while another is to establish a confirmation study. The power of a confirmation study must be high to ensure that the postulated quantitative trait loci can be verified. In the calculation of the experimental power, there are many issues that have to be addressed: size of the quantitative trait loci to be detected, significance level required, experimental design and expected heterozygosity for the design. To ensure marker assisted selection can be quickly implemented once quantitative trait loci are confirmed, DNA samples should be retained from daughters, and the sires and dams of elite sires.     

Accuracy of mapping quantitative trait loci in autogamous species

Summary The development of linkage maps with large numbers of molecular markers has stimulated the search for methods to map genes involved in quantitative traits (QTLs). A promising method, proposed by Lander and Botstein (1989), employs pairs of neighbouring markers to obtain maximum linkage information about the presence of a QTL within the enclosed chromosomal segment. In this paper the accuracy of this method was investigated by computer simulation. The results show that there is a reasonable probability of detecting QTLs that explain at least 5% of the total variance. For this purpose a minimum population of 200 backcross or F₂ individuals is necessary. Both the number of individuals and the relative size of the genotypic effect of the QTL are important factors determining the mapping precision. On the average, a QTL with 5% or 10% explained variance is mapped on an interval of 40 or 20 centiMorgans, respectively. Of course, QTLs with a larger genotypic effect will be located more precisely. It must be noted, however, that the interval length is rather variable.     

应用贝叶斯理论推断DNA分子标记基因型

引入贝叶斯理论用以从DNA分子标记的表现型（电泳谱带）推断其基因型（DNA来源）。结果表明,根据标记座位独立贫富而确定的遗传信息不完全标记的基因型概率,与根据邻近的遗传信息完全标记的基因型和有关重组率算得的相应贝叶斯概率,通常都有很大的差异,所以在进行数量性状基因定位和标记辅助选择等工作前前,应当计算每一个体基因组上所有遗传信息不完全座位的有关基因型的贝叶斯概率,文中列出计算未知基因型的贝叶斯概率的详细过程,也讨论了贝叶斯概率的若干推广应用。     

应用贝叶斯理论推断DNA分子标记基因型

引入贝叶斯理论用以从DNA分子标记的表现型（电泳谱带）推断其基因型（DNA来源）。结果表明,根据标记座位独立贫富而确定的遗传信息不完全标记的基因型概率,与根据令近的遗传信息完全标记的基因型和有关重组率算得的相应贝叶斯概率,通常都有很大的差异,所以在进行数量性状基因定位和标记辅助选择等工作之前,应当计算每一个体基因组上所有遗传信息不完全座位的有关基因型的贝叶斯概率。文中列出计算未知基因型的贝叶期概率的详细过程,也讨论了贝叶斯概率的若干推广应用。     

Power analysis of QTL detection in half-sib families using selective DNA pooling

Individual loci of economic importance (QTL) can be detected by comparing the inheritance of a trait and the inheritance of loci with alleles readily identifiable by laboratory methods (genetic markers). Data on allele segregation at the individual level are costly and alternatives have been proposed that make use of allele frequencies among progeny, rather than individual genotypes. Among the factors that may affect the power of the set up, the most important are those intrinsic to the QTL: the additive effect of the QTL, and its dominance, and distance between markers and QTL. Other factors are relative to the choice of animals and markers, such as the frequency of the QTL and marker alleles among dams and sires. Data collection may affect the detection power through the size of half-sib families, selection rate within families, and the technical error incurred when estimating genetic frequencies. We present results for a sensitivity analysis for QTL detection using pools of DNA from selected half-sibs. Simulations showed that conclusive detection may be achieved with families of at least 500 half-sibs if sires are chosen on the criteria that most of their marker alleles are either both missing, or one is fixed, among dams.     

数量性状基因座位及其在家禽中的定位

近年来随着现代分子生物学的发展 ,一些高效的分子遗传标记将各种畜禽的遗传图谱研究推向深入。为高密度、覆盖面广的连锁图谱的构建及QTL的定位奠定了基础。有关QTL的基本理论、QTL定位的步骤、QTL的检测方法以及家禽中定位的QTL已经展开了深入的研究 ,但目前QTL定位中存在诸多问题的问题