大豆抗倒伏性评价体系的建立及主要农艺性状QTL定位
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摘要
大豆起源于中国,是重要的食用植物蛋白和油脂来源,在我国国民经济发展以及世界粮食作物构成和国际油料作物生产中均占有重要地位。我国大豆常年种植面积约900万公顷,总产量约1600万吨,尚不能满足国内对大豆消费不断增长的需求,依靠进口国外转基因大豆。单产水平低和生产成本高是影响我国大豆生产发展的主要因素,提高大豆单产水平主要取决于大豆品种抗耐生物和非生物逆境性。
近年来灾害性天气频繁发生,严重制约了大豆等作物的丰产和稳产。我国南方春大豆生长中后期正值夏季多风雨时节,地表湿润,土壤疏松,易引发倒伏,对大豆产量和品质造成严重影响。因此开展抗倒性研究、培育高产抗倒伏品种,是应对灾害性天气、实现大豆高产稳产和优质的重要途径。
大豆倒伏性是多基因控制的数量性状,易受环境影响,仅以表型作为抗倒性育种的选择指标效率较低。本研究通过大豆品种资源田间鉴定试验,改进抗倒性鉴定和评价方法,并以中豆29和中豆32杂交衍生的重组自交系群体为材料,应用复合区间作图法开展大豆抗倒性及其相关形态性状和产量性状的遗传分析和QTL定位,旨在明确植株地上部和地下部性状与大豆抗倒性的关联,定位大豆抗倒性及主要农艺性状的数量性状位点(QTL),分析其遗传效应,为大豆抗倒性的分子标记辅助选择提供遗传学理论依据和分子标记。获得的主要结果如下:
1.通过抗倒性田间鉴测试验研究,提出综合植株地上部和地下部重量、茎秆强度、株高及分枝数等因子组成抗倒指数[(根重×茎秆强度)/(株高×茎叶重×分枝数)×100];确认结荚期抗倒指数与实际倒伏程度呈极显著负相关(r=-0.680),能够准确反映大豆种质的抗倒性差异,适合作为评价大豆种质抗倒伏能力的综合指标。
2.利用SSR标记构建了一张大豆遗传连锁图谱,该图谱包含28个连锁群,113个SSR标记,遗传距离1100.3 cM,平均标记间距为9.7 cM,最大距离32.4 cM,最小距离0.3 cM。与公共图谱比较,28个连锁群与公共图谱中的连锁群相对应,SSR标记在连锁群上的排列顺序与公共图谱基本一致。
3.对大豆倒伏性及其相关8个形态性状进行了QTL定位,共检测到45个QTL,分布于8个连锁群,可解释4.4%～50.1%的表型变异。其中位于F连锁群的倒伏主效QTL(qLD-15-1)和株高主效QTL(qPH-15-2),G连锁群的主茎节数OTL(qNMS-16-1),A2和L连锁群的根重QTL(qRW-1-2和qRW-24-1)在2个年份环境均重复检测到。在倒伏OTL的附近检测出株高、根重、茎叶重、茎粗、主茎节数和分枝数OTL,表明植株地上部和地下部性状与抗倒性普遍关联。大豆倒伏主效OTL的初定位为进一步开展精细定位和分子标记辅助育种提供了依据。
4.对大豆单株产量、产量构成因子等11个相关性状进行QTL检测和遗传效应分析,共检测到32个QTL,分布于6个连锁群,并主要集中于C2、F和I连锁群。控制百粒重、分枝荚数及有关产量性状的QTL与控制倒伏性QTL处于相同或相邻标记区间,表明产量相关性状与倒伏性存在一定的关联。I连锁群上的产量构成因子每荚粒数OTL和二、三、四粒荚数QTL不仅在2个年份均重复出现,解释的表型变异高(32%～65%),并且处于同一位置,每荚粒数和四粒荚数QTL与二、三粒荚数OTL的增效基因分别来自不同的亲本。这4个粒荚性状QTL的共位性与表型相关分析结果一致,证实每荚粒数和四粒荚数与二、三粒荚数分别由不同的机制调控,对于育种上探讨以改良大豆粒荚性状为途径提高大豆产量,提供了重要依据。
5.对不同发育阶段的大豆株高和茎粗同时进行非条件和条件QTL定位,在11个连锁群检测到株高和茎粗QTL均为13个。不同发育时期影响大豆株高和茎粗的OTL数量、加性效应和贡献率均不相同,QTL的表达具有时序性和选择性。株高和茎粗OTL在F连锁群上存在共位性,R1～R4期均有株高和茎粗QTL同时表达,但株高和茎粗OTL的增效基因不同。株高QTL表达次数多而茎粗QTL表达次数较少,前期(V4～R3)QTL表达数量多而后期(R4～R5)表达数量较少。株高和茎粗OTL的动态变化与表型相关分析结果一致,揭示了株高和茎粗在不同时段的相互关系和作用,对于适期选择秆粗抗倒的高产材料具有指导作用。
Soybean[Glycine max(L.) Merr.]originates in China.It is one of the important sources of protein and fat for human.In China,soybean is a major oilseed crop and is grown on about nine million hectares and total output about 16 million ton.Because of high velocity winds and frequent heavy rainfall,lodging is a common problem in production for soybean in the southern China.Lodging damage reduces both harvested yield and seed quality. Therefore,lodging resistance is one of the most important goals in soybean breeding.
Lodging represents a quantitative trait and is difficult to assess on a phenotypic basis. Marker-assisted selection(MAS) would become an important tool to improve lodging resistance in soybean.In this study,we mapped and characterised quantitative trait loci (QTLs) for lodging resistance,as well as morphological traits correlated with lodging and yield traits,in a segregating population of 165 recombinant inbred lines derived from the cross of the lodging-resistant soybean variety zhongdou 29 with the susceptible variety zhongdou 32.Field data were obtained in two years.The aim of current study was to explore genetics based on lodging resistance in soybean and gain useful molecular markers for molecular-assisted selection.The results are as following.
1.Index of lodging resistance(ILR) made up of different components was designed based on the mechanics as a comprehensive index to determine lodging resistance of 30 spring soybean cultivars.The ILR,(root weight×stem intension)/(plant height×shoot weight×branch number)×100,test at podding stage was significant negative relation with percentage of prostrate plant(r=-0.680).The test result could prove that the ILR might be used as an indicator to evaluate the lodging resistance of soybean objectively and accurately.
2.The linkage map including 28 linkage groups with 113 SSR markers.The map covered 1100.1 cM in length with an average marker interval of 9.7 cM,minimal interval of 0.3 cM and maximal interval of 32.4 cM.The order of markers in this map was identical to the order of "Consensus Linkage Map".
3.With the method of composite interval mapping(CIM) 45 QTL for lodging and related traits were detected on 8 linkage groups,explaining 4.4%-50.1%phenotypic variation. One QTL(qLD-15-1) of lodging and one QTL(qPH-15-2) of plant height on linkage group F were both detected in two years;one QTL of nodes on main stem and 2 QTLs of root weigh were detected in two years on linkage groups G,A2 and L respectively.The positions of some QTLs of plant height,root weight,shoot weight,stem diameter,nodes on main stem and branch number were close to those of lodging,showed that the lodging resistance is associated with the characters in aerial part and underground portion.The result might provide a basis for the further fine mapping and molecular marker-assisted breeding.
4.32 QTLs for yield,seed and pod traits were detected on 6 linkage groups,which majority were located on linkage groups C2,F and I.The QTLs of 100-grain weight and pod number on branch were in same or adjacent marker interval with the QTL of lodging, showed the correlation between yield-related traits and lodging.On linkage group I,the QTLs for yield components of number of seeds per pod,two-seed pod,three-seed pod,and four-seed pod were mapped to the same position in two years,explaining 32%-65% phenotypic variation.The positive alleles of QYLs for number of seeds per pod and four-seed pod,QTLs for two-seed pod and three-seed pod were from different parents,the four QTLs of seed and pod traits were mapped to the same loci.The result was consistent with the correlation analysis of phenotypic traits.The result showed that the regulatory mechanisms of number of seeds per pod and four-seed pod,two-seed pod and three-seed pod were different,which may provide an important basis for promotion of soybean yield by improving seed and pod traits.
5.Based on unconditional and conditional QTL mapping methods,13 QTLs for plant height and 13 QTLs for stem diameter were detected on eleven linkage groups.The number, additive effects and contribution rate of QTLs for plant height and stem diameter were different at different measuring stages,which indicated that these QTLs expressed timely and electively.The QTLs of plant height and stem diameter on linkage groups F were mapped to the same loci and they were expressed simultaneously from R1 to R4 stages.In the early stages QTLs expressed more than in late stages,and QTLs for plant height expressed more times than QTLs for stem diameter.The positive alleles of QTLs for plant height and stem diameter were from different parents.The developmental dynamic of QTLs for plant height and stem diameter was consistent with the correlation analysis ofphenotypic traits at different developmental stages.
近年来灾害性天气频繁发生,严重制约了大豆等作物的丰产和稳产。我国南方春大豆生长中后期正值夏季多风雨时节,地表湿润,土壤疏松,易引发倒伏,对大豆产量和品质造成严重影响。因此开展抗倒性研究、培育高产抗倒伏品种,是应对灾害性天气、实现大豆高产稳产和优质的重要途径。
大豆倒伏性是多基因控制的数量性状,易受环境影响,仅以表型作为抗倒性育种的选择指标效率较低。本研究通过大豆品种资源田间鉴定试验,改进抗倒性鉴定和评价方法,并以中豆29和中豆32杂交衍生的重组自交系群体为材料,应用复合区间作图法开展大豆抗倒性及其相关形态性状和产量性状的遗传分析和QTL定位,旨在明确植株地上部和地下部性状与大豆抗倒性的关联,定位大豆抗倒性及主要农艺性状的数量性状位点(QTL),分析其遗传效应,为大豆抗倒性的分子标记辅助选择提供遗传学理论依据和分子标记。获得的主要结果如下:
1.通过抗倒性田间鉴测试验研究,提出综合植株地上部和地下部重量、茎秆强度、株高及分枝数等因子组成抗倒指数[(根重×茎秆强度)/(株高×茎叶重×分枝数)×100];确认结荚期抗倒指数与实际倒伏程度呈极显著负相关(r=-0.680),能够准确反映大豆种质的抗倒性差异,适合作为评价大豆种质抗倒伏能力的综合指标。
2.利用SSR标记构建了一张大豆遗传连锁图谱,该图谱包含28个连锁群,113个SSR标记,遗传距离1100.3 cM,平均标记间距为9.7 cM,最大距离32.4 cM,最小距离0.3 cM。与公共图谱比较,28个连锁群与公共图谱中的连锁群相对应,SSR标记在连锁群上的排列顺序与公共图谱基本一致。
3.对大豆倒伏性及其相关8个形态性状进行了QTL定位,共检测到45个QTL,分布于8个连锁群,可解释4.4%～50.1%的表型变异。其中位于F连锁群的倒伏主效QTL(qLD-15-1)和株高主效QTL(qPH-15-2),G连锁群的主茎节数OTL(qNMS-16-1),A2和L连锁群的根重QTL(qRW-1-2和qRW-24-1)在2个年份环境均重复检测到。在倒伏OTL的附近检测出株高、根重、茎叶重、茎粗、主茎节数和分枝数OTL,表明植株地上部和地下部性状与抗倒性普遍关联。大豆倒伏主效OTL的初定位为进一步开展精细定位和分子标记辅助育种提供了依据。
4.对大豆单株产量、产量构成因子等11个相关性状进行QTL检测和遗传效应分析,共检测到32个QTL,分布于6个连锁群,并主要集中于C2、F和I连锁群。控制百粒重、分枝荚数及有关产量性状的QTL与控制倒伏性QTL处于相同或相邻标记区间,表明产量相关性状与倒伏性存在一定的关联。I连锁群上的产量构成因子每荚粒数OTL和二、三、四粒荚数QTL不仅在2个年份均重复出现,解释的表型变异高(32%～65%),并且处于同一位置,每荚粒数和四粒荚数QTL与二、三粒荚数OTL的增效基因分别来自不同的亲本。这4个粒荚性状QTL的共位性与表型相关分析结果一致,证实每荚粒数和四粒荚数与二、三粒荚数分别由不同的机制调控,对于育种上探讨以改良大豆粒荚性状为途径提高大豆产量,提供了重要依据。
5.对不同发育阶段的大豆株高和茎粗同时进行非条件和条件QTL定位,在11个连锁群检测到株高和茎粗QTL均为13个。不同发育时期影响大豆株高和茎粗的OTL数量、加性效应和贡献率均不相同,QTL的表达具有时序性和选择性。株高和茎粗OTL在F连锁群上存在共位性,R1～R4期均有株高和茎粗QTL同时表达,但株高和茎粗OTL的增效基因不同。株高QTL表达次数多而茎粗QTL表达次数较少,前期(V4～R3)QTL表达数量多而后期(R4～R5)表达数量较少。株高和茎粗OTL的动态变化与表型相关分析结果一致,揭示了株高和茎粗在不同时段的相互关系和作用,对于适期选择秆粗抗倒的高产材料具有指导作用。
Soybean[Glycine max(L.) Merr.]originates in China.It is one of the important sources of protein and fat for human.In China,soybean is a major oilseed crop and is grown on about nine million hectares and total output about 16 million ton.Because of high velocity winds and frequent heavy rainfall,lodging is a common problem in production for soybean in the southern China.Lodging damage reduces both harvested yield and seed quality. Therefore,lodging resistance is one of the most important goals in soybean breeding.
Lodging represents a quantitative trait and is difficult to assess on a phenotypic basis. Marker-assisted selection(MAS) would become an important tool to improve lodging resistance in soybean.In this study,we mapped and characterised quantitative trait loci (QTLs) for lodging resistance,as well as morphological traits correlated with lodging and yield traits,in a segregating population of 165 recombinant inbred lines derived from the cross of the lodging-resistant soybean variety zhongdou 29 with the susceptible variety zhongdou 32.Field data were obtained in two years.The aim of current study was to explore genetics based on lodging resistance in soybean and gain useful molecular markers for molecular-assisted selection.The results are as following.
1.Index of lodging resistance(ILR) made up of different components was designed based on the mechanics as a comprehensive index to determine lodging resistance of 30 spring soybean cultivars.The ILR,(root weight×stem intension)/(plant height×shoot weight×branch number)×100,test at podding stage was significant negative relation with percentage of prostrate plant(r=-0.680).The test result could prove that the ILR might be used as an indicator to evaluate the lodging resistance of soybean objectively and accurately.
2.The linkage map including 28 linkage groups with 113 SSR markers.The map covered 1100.1 cM in length with an average marker interval of 9.7 cM,minimal interval of 0.3 cM and maximal interval of 32.4 cM.The order of markers in this map was identical to the order of "Consensus Linkage Map".
3.With the method of composite interval mapping(CIM) 45 QTL for lodging and related traits were detected on 8 linkage groups,explaining 4.4%-50.1%phenotypic variation. One QTL(qLD-15-1) of lodging and one QTL(qPH-15-2) of plant height on linkage group F were both detected in two years;one QTL of nodes on main stem and 2 QTLs of root weigh were detected in two years on linkage groups G,A2 and L respectively.The positions of some QTLs of plant height,root weight,shoot weight,stem diameter,nodes on main stem and branch number were close to those of lodging,showed that the lodging resistance is associated with the characters in aerial part and underground portion.The result might provide a basis for the further fine mapping and molecular marker-assisted breeding.
4.32 QTLs for yield,seed and pod traits were detected on 6 linkage groups,which majority were located on linkage groups C2,F and I.The QTLs of 100-grain weight and pod number on branch were in same or adjacent marker interval with the QTL of lodging, showed the correlation between yield-related traits and lodging.On linkage group I,the QTLs for yield components of number of seeds per pod,two-seed pod,three-seed pod,and four-seed pod were mapped to the same position in two years,explaining 32%-65% phenotypic variation.The positive alleles of QYLs for number of seeds per pod and four-seed pod,QTLs for two-seed pod and three-seed pod were from different parents,the four QTLs of seed and pod traits were mapped to the same loci.The result was consistent with the correlation analysis of phenotypic traits.The result showed that the regulatory mechanisms of number of seeds per pod and four-seed pod,two-seed pod and three-seed pod were different,which may provide an important basis for promotion of soybean yield by improving seed and pod traits.
5.Based on unconditional and conditional QTL mapping methods,13 QTLs for plant height and 13 QTLs for stem diameter were detected on eleven linkage groups.The number, additive effects and contribution rate of QTLs for plant height and stem diameter were different at different measuring stages,which indicated that these QTLs expressed timely and electively.The QTLs of plant height and stem diameter on linkage groups F were mapped to the same loci and they were expressed simultaneously from R1 to R4 stages.In the early stages QTLs expressed more than in late stages,and QTLs for plant height expressed more times than QTLs for stem diameter.The positive alleles of QTLs for plant height and stem diameter were from different parents.The developmental dynamic of QTLs for plant height and stem diameter was consistent with the correlation analysis ofphenotypic traits at different developmental stages.
引文
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