小麦骨干亲本矮孟牛姊妹系基因组差异及1RS遗传效应分析
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摘要
小麦骨干亲本矮孟牛在我国品种的遗传改良中发挥了重要作用，特别是矮孟牛V型不仅曾是审定的大面积推广品种，而且以其作为杂交亲本选育出多个优良小麦品种和一批优良种质系。本研究通过表型和基因组扫描分析，阐明小麦骨干亲本矮孟牛（V型）的遗传本质，为探讨小麦骨干亲本的成因及其应用、新型骨干亲本的创制提供理论指导。同时，通过构建不同的小麦遗传群体，分析明确1RS对产量和品质的遗传效应，为其在小麦遗传改良中的利用提供依据。
通过研究获得以下主要结果：
（1）利用1B和1R分子标记和基因组原位杂交的方法对“矮孟牛”7个姊妹系的遗传差异进行了鉴定。基因组原位杂交结果证明，在矮孟牛Ⅱ型、Ⅳ型-Ⅶ型中含有黑麦的杂交信号，而在矮孟牛Ⅰ型和Ⅲ型中不含黑麦的杂交信号。PCR结果显示，矮孟牛Ⅱ型、Ⅳ-Ⅶ型中含有1RS和1BL，而不含1BS和1RL；矮孟牛Ⅰ型和Ⅲ型中含有正常的1B染色体。
（2）采用田间试验鉴定和基因组分子标记分析相结合的方法，对冬小麦种质“矮孟牛”7个姊妹系的性状和基因组差异比较分析的结果表明，7个类型在调查的株高、穗长、穗粒数、粒长、粒宽、单株穗数等16个性状方面存在差异，其中矮孟牛V型的主要产量构成因素等性状比较协调，总体优于其他6个类型。利用656对基因组分子标记对矮孟牛7个类型进行全基因组遗传差异分析，并利用GGT2.0绘制了7个类型的遗传差异图谱。在矮孟牛V型的1A、1B、2D、3A、4D、7A染色体上检测到8个特异位点。利用矮丰3号/牛朱特、孟县201/牛朱特、矮丰3号//孟县201/牛朱特、孟县201//矮丰3号/牛朱特，矮孟牛V型/济麦22和V型/泰农18的F2分离群体，经IciMappingV3.0单标记QTL作图分析，发现矮孟牛V型部分特异位点附近存在与产量构成因素等重要性状相关的QTL。矮孟牛V型的基因组特异位点可能是它作为骨干亲本区别于其他姊妹系的重要基因组特征。
（3）利用济麦22/矮孟牛V型和泰农18/矮孟牛V型的F2分离群体进行单标记QTL分析，发现V型的1RS染色体上存在与增加小穗数/穗、单穗粒重、可育小穗数/穗和百粒重相关的QTL。矮孟牛V型中1RS的存在可能是其成为骨干亲本的重要原因之一。同时，利用1RS对潍麦8号（1B/1R易位系）/济麦20和潍麦8号/烟农19的RIL群体进行单标记QTL分析，比较了4个不同群体中1RS对产量和品质相关性状的遗传效应。结果表明，不同群体中1RS对产量相关性状的遗传效应是不同的（开花期除外），而对品质相关性状的遗传效应是相同的。
（4）从潍麦8号与济麦20的RIL群体中选取10个1BL/1RS纯合易位系和10个1B株系，通过田间试验和环境与基因型两因素方差分析结果表明，1BL/1RS易位系的籽粒产量、地上部生物产量、株高、穗粒数和小穗数高于1B株系，而其千粒重低于1B株系；1BL/1RS易位系的品质性状较差，它与1B株系相比具有较低的籽粒硬度、容重、沉降值和面团延展性。
Backbone parents Aimengniu (AMN), especially AMN-derived type V, play animportant role in the renewal of Chinese wheat varieties. AMN-derived type V was anauthorized variety spraying for large areas, from which many wheat varieties and germplasmswith good agronomic characters were selected. In the present study, phenotypic and genomicanalysis of Aimengniu were conducted, with an aim to elucidate the genetic basis of backboneparent AMN-derived type V. These results will provid the theory guidance for the creating ofnew backbone parent. The genetic effect of1RS on yield and quality in different geneticbackgrounds were also specified, which provided the genetic basis for the utilization of1RSin wheat genetic improvement. The main results were as follows:
(1) Molecular markers of1B and1R as well as genomic in situ hybridization (GISH)were used to identify genetic differences in the AMN-derived seven siblines. The results ofGISH showed the presence of rye signal in AMN-derived type II and IV to VII, while absencein AMN-derived type I and III. PCR results of primers specific for1R and1B proved thepresence of both1RS and1BL chromatin and absence of1BS and1RL in AMN-derived typeII and IV to VII, while AMN-derived type I and III contained common1B chromatin. Theseresults were well agreed with that of GISH.
(2) The genetic and phenotypic differences among the AMN-derived seven siblines werecompared based on656molecular markers and16agronomic traits including plant height,panicle length, grain number per panicle, grain length, grain width, panicle number per plant.There were obvious variations among the AMN-derived seven siblines in the traits surveyed.The AMN-derive type V showed better performance on yield components than other types. Agraphical genotype integrated map was constructed by GGT2.0using the molecular markerdata, which showed the inheritance of fragments from the three parents to AMN. Six F2populations (Aifeng3/Neuzucht, Mengxian201/Neuzucht, Aifeng3//Mengxian201/Neuzucht, Mengxian201//Aifeng3/Neuzucht, AMN V/Jimai22and AMN V/Tainong18)were also developed to detect quantitative trait loci (QTLs) for yield related traits in theAMN-derived type V. Eight AMN-derived type V specific loci were detected onchromosomes1A,1B,2D,3A,4D, and7A, which were associated with QTLs for yield components. These specific loci might discriminate the type V from the remaining types atthe genomic level.
(3) F2populations of AMN V/Jimai22and AMN V/Tainong18were used to detect yieldrelated quantitative trait loci (QTLs) in1RS of AMN-derived type V. The results showed that1RS was associated with QTLs for increasing spikelet number per spike, grain weight perspike, fertile spikelet number per spike and100-kernel weight. The1RS chromosome ofAMN-derived type V might provide the information for understanding the genetic basis ofAMN-derive type V as a backbone parent in wheat breeding. At the same time, RILs ofWeimai8/Jimai20and Weimai8/Yannong19were executed single marker analysis based on1RS-specific markers. The results of the genetic effect of1RS on the4populations showedthat1RS had different genetic effect on yield related traits (excepted for flower date), while ithad the same effect on quality related traits.
(4) Ten1BL/1RS translocation lines and101BS lines selected from the RILs ofWeimai8/Jimai20were excuted two-way ANOVA of environment and chromosome classes.The results showed that1BL/1RS had a higher grain yield, above-ground biomass, plantheight, kernel number per spike, and spikelet number per spike. However, the1000-grainweight was lower. In contrast to the superior grain yield potential, the presence of1BL/1RShad a significant negative effect on the quality, i.e., the kernel hardness, testweight, thesedimentation volume and tractility were lower.
通过研究获得以下主要结果：
（1）利用1B和1R分子标记和基因组原位杂交的方法对“矮孟牛”7个姊妹系的遗传差异进行了鉴定。基因组原位杂交结果证明，在矮孟牛Ⅱ型、Ⅳ型-Ⅶ型中含有黑麦的杂交信号，而在矮孟牛Ⅰ型和Ⅲ型中不含黑麦的杂交信号。PCR结果显示，矮孟牛Ⅱ型、Ⅳ-Ⅶ型中含有1RS和1BL，而不含1BS和1RL；矮孟牛Ⅰ型和Ⅲ型中含有正常的1B染色体。
（2）采用田间试验鉴定和基因组分子标记分析相结合的方法，对冬小麦种质“矮孟牛”7个姊妹系的性状和基因组差异比较分析的结果表明，7个类型在调查的株高、穗长、穗粒数、粒长、粒宽、单株穗数等16个性状方面存在差异，其中矮孟牛V型的主要产量构成因素等性状比较协调，总体优于其他6个类型。利用656对基因组分子标记对矮孟牛7个类型进行全基因组遗传差异分析，并利用GGT2.0绘制了7个类型的遗传差异图谱。在矮孟牛V型的1A、1B、2D、3A、4D、7A染色体上检测到8个特异位点。利用矮丰3号/牛朱特、孟县201/牛朱特、矮丰3号//孟县201/牛朱特、孟县201//矮丰3号/牛朱特，矮孟牛V型/济麦22和V型/泰农18的F2分离群体，经IciMappingV3.0单标记QTL作图分析，发现矮孟牛V型部分特异位点附近存在与产量构成因素等重要性状相关的QTL。矮孟牛V型的基因组特异位点可能是它作为骨干亲本区别于其他姊妹系的重要基因组特征。
（3）利用济麦22/矮孟牛V型和泰农18/矮孟牛V型的F2分离群体进行单标记QTL分析，发现V型的1RS染色体上存在与增加小穗数/穗、单穗粒重、可育小穗数/穗和百粒重相关的QTL。矮孟牛V型中1RS的存在可能是其成为骨干亲本的重要原因之一。同时，利用1RS对潍麦8号（1B/1R易位系）/济麦20和潍麦8号/烟农19的RIL群体进行单标记QTL分析，比较了4个不同群体中1RS对产量和品质相关性状的遗传效应。结果表明，不同群体中1RS对产量相关性状的遗传效应是不同的（开花期除外），而对品质相关性状的遗传效应是相同的。
（4）从潍麦8号与济麦20的RIL群体中选取10个1BL/1RS纯合易位系和10个1B株系，通过田间试验和环境与基因型两因素方差分析结果表明，1BL/1RS易位系的籽粒产量、地上部生物产量、株高、穗粒数和小穗数高于1B株系，而其千粒重低于1B株系；1BL/1RS易位系的品质性状较差，它与1B株系相比具有较低的籽粒硬度、容重、沉降值和面团延展性。
Backbone parents Aimengniu (AMN), especially AMN-derived type V, play animportant role in the renewal of Chinese wheat varieties. AMN-derived type V was anauthorized variety spraying for large areas, from which many wheat varieties and germplasmswith good agronomic characters were selected. In the present study, phenotypic and genomicanalysis of Aimengniu were conducted, with an aim to elucidate the genetic basis of backboneparent AMN-derived type V. These results will provid the theory guidance for the creating ofnew backbone parent. The genetic effect of1RS on yield and quality in different geneticbackgrounds were also specified, which provided the genetic basis for the utilization of1RSin wheat genetic improvement. The main results were as follows:
(1) Molecular markers of1B and1R as well as genomic in situ hybridization (GISH)were used to identify genetic differences in the AMN-derived seven siblines. The results ofGISH showed the presence of rye signal in AMN-derived type II and IV to VII, while absencein AMN-derived type I and III. PCR results of primers specific for1R and1B proved thepresence of both1RS and1BL chromatin and absence of1BS and1RL in AMN-derived typeII and IV to VII, while AMN-derived type I and III contained common1B chromatin. Theseresults were well agreed with that of GISH.
(2) The genetic and phenotypic differences among the AMN-derived seven siblines werecompared based on656molecular markers and16agronomic traits including plant height,panicle length, grain number per panicle, grain length, grain width, panicle number per plant.There were obvious variations among the AMN-derived seven siblines in the traits surveyed.The AMN-derive type V showed better performance on yield components than other types. Agraphical genotype integrated map was constructed by GGT2.0using the molecular markerdata, which showed the inheritance of fragments from the three parents to AMN. Six F2populations (Aifeng3/Neuzucht, Mengxian201/Neuzucht, Aifeng3//Mengxian201/Neuzucht, Mengxian201//Aifeng3/Neuzucht, AMN V/Jimai22and AMN V/Tainong18)were also developed to detect quantitative trait loci (QTLs) for yield related traits in theAMN-derived type V. Eight AMN-derived type V specific loci were detected onchromosomes1A,1B,2D,3A,4D, and7A, which were associated with QTLs for yield components. These specific loci might discriminate the type V from the remaining types atthe genomic level.
(3) F2populations of AMN V/Jimai22and AMN V/Tainong18were used to detect yieldrelated quantitative trait loci (QTLs) in1RS of AMN-derived type V. The results showed that1RS was associated with QTLs for increasing spikelet number per spike, grain weight perspike, fertile spikelet number per spike and100-kernel weight. The1RS chromosome ofAMN-derived type V might provide the information for understanding the genetic basis ofAMN-derive type V as a backbone parent in wheat breeding. At the same time, RILs ofWeimai8/Jimai20and Weimai8/Yannong19were executed single marker analysis based on1RS-specific markers. The results of the genetic effect of1RS on the4populations showedthat1RS had different genetic effect on yield related traits (excepted for flower date), while ithad the same effect on quality related traits.
(4) Ten1BL/1RS translocation lines and101BS lines selected from the RILs ofWeimai8/Jimai20were excuted two-way ANOVA of environment and chromosome classes.The results showed that1BL/1RS had a higher grain yield, above-ground biomass, plantheight, kernel number per spike, and spikelet number per spike. However, the1000-grainweight was lower. In contrast to the superior grain yield potential, the presence of1BL/1RShad a significant negative effect on the quality, i.e., the kernel hardness, testweight, thesedimentation volume and tractility were lower.
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