棉花海陆杂交—回交后代群体纤维长度性状的遗传分析
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摘要
本研究采用DNA分子标记技术（如SSR、SRAP等），以新疆“自育”的海岛棉和陆地棉为亲本及其衍生的F2群体等为材料，绘制结合分子标记的遗传连锁图谱。结合考查的F_2和F_(2:3)群体的农艺性状及纤维特性，在已构建的遗传连锁图谱上分析数量性状基因位点、检测并得到相应的QTL；利用种间杂交-回交高世代群体（BC_4F_2/BC_4F_(2:3)）剖析数量性状基因位点遗传变异的分子机制，旨在寻找到与棉花纤维长度性状相关的遗传基因位点，为分子标记辅助和分子水平上的遗传改良提供依据。获得了如下的结果。
1.对丰产性优良、抗性强的系9和纤维品质性状优良的新海16及其所产生的后代群体进行比较分析。结果表明，系9和新海16的纤维长度性状的遗传特性符合杂交育种的亲本选配原则；且纤维长度性状呈现正态分布，能进行QTL检测。
2.以陆地棉和海岛棉为材料，利用本实验建立和优化的分子标记体系，对48份棉花资源进行分析和基于本实验新开发的棉花纤维序列的93对EST-SSR适用性检测。结果表明，系9具有可以作为陆地棉基础种质的特性；新海16具有海岛棉的代表性。本实验优化的分子标记技术体系是可行的。
3.以系9作母本，新海16作父本，构建了F2分离群体（297个单株），结合SSRs,SRAPs和EST-SSRs技术，构建连锁图谱。用800多对SSR引物，93对EST-SSR引物、300多个SRAP引物组合分析系9和新海16间的多态性，获得了245对(占67.1%)多态性SSR引物，EST-SSR引物88条(占24.1%)及32对(占8.8%)SRAP引物的组合。
4.利用有多态性的引物对F_2群体扩增，结合QTL分析，获得了229对SSR引物，29个SRAP引物组合，88对EST-SSR引物。用QTL-IciMapping对多态性标记进行连锁分析。得到一张含346个标记的22个连锁群。图谱总长为4307.73cM，占棉花总基因组（5000cM）的86.15%；平均标记间距为12.45cM。
5.对F_2群体的每个植株及其衍生的F_2：3家系(设置两重复)的纤维长度性状进行考查，用完备区间作图法进行QTL检测，LOD域值为3.0，共定位了34个QTLs：分析得到在第四连锁群上检测到的QTL有两个显著标记，解释表型变异分别为17.3072%（NAU3207/me4em5）和13.7922%(me3em10/B3171)
6.以系9作母本，纤维品质优良的海岛棉新海16作父本，杂交后回交4代，通过单籽粒传法最终构建了一个种间杂交-回交高世代的渗入系群体，含有234个BC_4F_(2:3)株系。
7.田间种植BC4F2:3家系，随机区组三次重复，考查纤维品质性状，各性状的频率分布显示:纤维长度性状频率分布图的峰偏向新海16；相关性分析表明：大多数性状之间均存在差异显著，但相关关系不等。以利用系9与新海16杂交的F2代所构建的连锁图谱的引物为参照，结合查询的与纤维长度有关的标记，用相关引物对BC4F2群体进行分析，用QTL-IciMapping对利用的标记产生的数据实施遗传作图。绘制了一张19个连锁群，包含278个标记。其总长为4351.65cM，占棉花总基因组（5000cM）的87.03%；标记间平均间距为15.65cM。
8.利用标记数据与性状数据进行QTL检测分析，共检测到74个纤维长度性状相关的QTL，检测到的QTL有7个显著标记，解释表型变异在9.1078%和13.6005%之间。且都为显性增益占主导，说明海陆杂交后代的纤维长度性状具有较强的优势，应在育种中加强应用。
In this study, genetic linkage maps for tetraploid cotton were constructed by means of DNA-basedmarkers based on two segregation populations. Cotton agronomic trait and fiber quality were investigatedin the population of F_2individuals and corresponding F_(2:3)families and High backcross generations(BC_4F_2and BC_4F_(2:3)). On the basis of genetic map constructed; quantitative traits analysis and QTLs detection wasperformed. The purpose of this study was to find the genetic loci associated significantly with fiber length,so the further research and application (such as marker-aided selection, map-based cloning) could beconducted. The results obtained in this study were as follows.
1. The high yielding fine, strong resistance of xi9and good fiber quality traits of xinhai16and itsoffspring produced by group a comparative study. Results showed that fiber length between xi9andxinhai16is fit for cross breeding parent matching principle; And fiber length properties presented normaldistribution, can detect QTL.
2. Upland cotton and island cotton as material, the use of the experimental set up and optimizedmolecular marker system, analysis of48cotton resources and based on this experiment is a newdevelopment of cotton fiber series of applicability of93EST-SSR detection, the results show that thesystem can have as a foundation for upland cotton germplasm characteristics; New sea16representativewith sea island cotton. This experiment optimized molecular marker technology system is feasible.
3.An interspecific F2population consisted of297plants, which was developed from the cross betweenXi9(characterized as high fiber yield) and Xinhai16(characterized as excellent fiber quality), wasgenotyped with SSR, SRAP and EST-SSR markers. Over1200(pairs of) primers (combinations) were usedto survey the polymorphism between the parents, and the result showed that245pairs of SSR primers(67.1%),88EST-SSR primers (24.1.3%),32SRAP primer combinations (8.8%) could amplifypolymorphic bands.
4.The marker data were subsequently analyzed using QTL-IcIMapping2.3. Among the229pairs ofSSR primers,88EST-SSR primers,29SRAP primer were arranged into22linkage groups. The linkagemap covered4307.73cM with an average distance of12.45cM between two markers.
5.The F_2plants and corresponding F_(2:3)families (arranged in2replicates in a random block design)were used for scoring the phenotypes of fiber length. The present genetic linkage map and traits data wasused to identify and map the quantitative trait loci (QTLs) affecting fiber length traits in297F_(2:3)familylines. The method of composite interval mapping was applied to search for QTLs by QTL-ICIMappingV2.3. The log odds-ratio threshold of QTL detection was3.0, on the basis of which this analysis yielded34QTLs: two QTLs controlling fiber length are significant marks in linkage4th,explained17.30729%(NAU3207/me4em5）and13.7922%(me3em10/B3171) of phenotypic variation (PVE).
6.An advanced-generation population was developed by first crossing G. hirsutum cv.xi9and G.barbadense cv. Xinhai16(characterized as excellent fiber quality),then single seed descent to BC_4F_2generation. This population consisted of234BC_4F_2lines, which containing chromatin segmentintrogressived from xinhai16to xi9background.
7.Field experiment was carried out for234BC_4F_(2:3)families in2009(arranged in3replicates in arandom block design), and the phenotypes of fiber quality were scored. The phenotypic distribution of theindividual introgression lines for five traits indicated the trend of regression of individual lines to xinhai16and the wide variance for each trait. The result from correlation analysis among the five traits showed that:trait combinations were significantly correlated except the combinations of micronaire and elongation, butthe coefficients were different.Following the order and primers of a linkage map developed from the F_2populationfrom the same cross, several hundred of SSR markers were used in this study to detect genotypes.278primers were arranged into19linkage groups. The linkage map covered4351.65cM with an averagedistance of15.65cM between two markers.
8.The method of composite interval mapping was applied to search for QTLs by QTL-ICIMappingV2.3. The log odds-ratio threshold of QTL detection was3.0, on the basis of which this analysis yielded74QTLs: seven QTLs controlling fiber length are significant marks in linkage all,explained from9.1078%to13.6005%of phenotypic variation (PVE).And there are the dominant gain dominant, That said thesea-land fiber length of the hybrid offspring traits has a strong advantage, enhancing the use in breeding
1.对丰产性优良、抗性强的系9和纤维品质性状优良的新海16及其所产生的后代群体进行比较分析。结果表明，系9和新海16的纤维长度性状的遗传特性符合杂交育种的亲本选配原则；且纤维长度性状呈现正态分布，能进行QTL检测。
2.以陆地棉和海岛棉为材料，利用本实验建立和优化的分子标记体系，对48份棉花资源进行分析和基于本实验新开发的棉花纤维序列的93对EST-SSR适用性检测。结果表明，系9具有可以作为陆地棉基础种质的特性；新海16具有海岛棉的代表性。本实验优化的分子标记技术体系是可行的。
3.以系9作母本，新海16作父本，构建了F2分离群体（297个单株），结合SSRs,SRAPs和EST-SSRs技术，构建连锁图谱。用800多对SSR引物，93对EST-SSR引物、300多个SRAP引物组合分析系9和新海16间的多态性，获得了245对(占67.1%)多态性SSR引物，EST-SSR引物88条(占24.1%)及32对(占8.8%)SRAP引物的组合。
4.利用有多态性的引物对F_2群体扩增，结合QTL分析，获得了229对SSR引物，29个SRAP引物组合，88对EST-SSR引物。用QTL-IciMapping对多态性标记进行连锁分析。得到一张含346个标记的22个连锁群。图谱总长为4307.73cM，占棉花总基因组（5000cM）的86.15%；平均标记间距为12.45cM。
5.对F_2群体的每个植株及其衍生的F_2：3家系(设置两重复)的纤维长度性状进行考查，用完备区间作图法进行QTL检测，LOD域值为3.0，共定位了34个QTLs：分析得到在第四连锁群上检测到的QTL有两个显著标记，解释表型变异分别为17.3072%（NAU3207/me4em5）和13.7922%(me3em10/B3171)
6.以系9作母本，纤维品质优良的海岛棉新海16作父本，杂交后回交4代，通过单籽粒传法最终构建了一个种间杂交-回交高世代的渗入系群体，含有234个BC_4F_(2:3)株系。
7.田间种植BC4F2:3家系，随机区组三次重复，考查纤维品质性状，各性状的频率分布显示:纤维长度性状频率分布图的峰偏向新海16；相关性分析表明：大多数性状之间均存在差异显著，但相关关系不等。以利用系9与新海16杂交的F2代所构建的连锁图谱的引物为参照，结合查询的与纤维长度有关的标记，用相关引物对BC4F2群体进行分析，用QTL-IciMapping对利用的标记产生的数据实施遗传作图。绘制了一张19个连锁群，包含278个标记。其总长为4351.65cM，占棉花总基因组（5000cM）的87.03%；标记间平均间距为15.65cM。
8.利用标记数据与性状数据进行QTL检测分析，共检测到74个纤维长度性状相关的QTL，检测到的QTL有7个显著标记，解释表型变异在9.1078%和13.6005%之间。且都为显性增益占主导，说明海陆杂交后代的纤维长度性状具有较强的优势，应在育种中加强应用。
In this study, genetic linkage maps for tetraploid cotton were constructed by means of DNA-basedmarkers based on two segregation populations. Cotton agronomic trait and fiber quality were investigatedin the population of F_2individuals and corresponding F_(2:3)families and High backcross generations(BC_4F_2and BC_4F_(2:3)). On the basis of genetic map constructed; quantitative traits analysis and QTLs detection wasperformed. The purpose of this study was to find the genetic loci associated significantly with fiber length,so the further research and application (such as marker-aided selection, map-based cloning) could beconducted. The results obtained in this study were as follows.
1. The high yielding fine, strong resistance of xi9and good fiber quality traits of xinhai16and itsoffspring produced by group a comparative study. Results showed that fiber length between xi9andxinhai16is fit for cross breeding parent matching principle; And fiber length properties presented normaldistribution, can detect QTL.
2. Upland cotton and island cotton as material, the use of the experimental set up and optimizedmolecular marker system, analysis of48cotton resources and based on this experiment is a newdevelopment of cotton fiber series of applicability of93EST-SSR detection, the results show that thesystem can have as a foundation for upland cotton germplasm characteristics; New sea16representativewith sea island cotton. This experiment optimized molecular marker technology system is feasible.
3.An interspecific F2population consisted of297plants, which was developed from the cross betweenXi9(characterized as high fiber yield) and Xinhai16(characterized as excellent fiber quality), wasgenotyped with SSR, SRAP and EST-SSR markers. Over1200(pairs of) primers (combinations) were usedto survey the polymorphism between the parents, and the result showed that245pairs of SSR primers(67.1%),88EST-SSR primers (24.1.3%),32SRAP primer combinations (8.8%) could amplifypolymorphic bands.
4.The marker data were subsequently analyzed using QTL-IcIMapping2.3. Among the229pairs ofSSR primers,88EST-SSR primers,29SRAP primer were arranged into22linkage groups. The linkagemap covered4307.73cM with an average distance of12.45cM between two markers.
5.The F_2plants and corresponding F_(2:3)families (arranged in2replicates in a random block design)were used for scoring the phenotypes of fiber length. The present genetic linkage map and traits data wasused to identify and map the quantitative trait loci (QTLs) affecting fiber length traits in297F_(2:3)familylines. The method of composite interval mapping was applied to search for QTLs by QTL-ICIMappingV2.3. The log odds-ratio threshold of QTL detection was3.0, on the basis of which this analysis yielded34QTLs: two QTLs controlling fiber length are significant marks in linkage4th,explained17.30729%(NAU3207/me4em5）and13.7922%(me3em10/B3171) of phenotypic variation (PVE).
6.An advanced-generation population was developed by first crossing G. hirsutum cv.xi9and G.barbadense cv. Xinhai16(characterized as excellent fiber quality),then single seed descent to BC_4F_2generation. This population consisted of234BC_4F_2lines, which containing chromatin segmentintrogressived from xinhai16to xi9background.
7.Field experiment was carried out for234BC_4F_(2:3)families in2009(arranged in3replicates in arandom block design), and the phenotypes of fiber quality were scored. The phenotypic distribution of theindividual introgression lines for five traits indicated the trend of regression of individual lines to xinhai16and the wide variance for each trait. The result from correlation analysis among the five traits showed that:trait combinations were significantly correlated except the combinations of micronaire and elongation, butthe coefficients were different.Following the order and primers of a linkage map developed from the F_2populationfrom the same cross, several hundred of SSR markers were used in this study to detect genotypes.278primers were arranged into19linkage groups. The linkage map covered4351.65cM with an averagedistance of15.65cM between two markers.
8.The method of composite interval mapping was applied to search for QTLs by QTL-ICIMappingV2.3. The log odds-ratio threshold of QTL detection was3.0, on the basis of which this analysis yielded74QTLs: seven QTLs controlling fiber length are significant marks in linkage all,explained from9.1078%to13.6005%of phenotypic variation (PVE).And there are the dominant gain dominant, That said thesea-land fiber length of the hybrid offspring traits has a strong advantage, enhancing the use in breeding
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