大白菜分子遗传图谱构建及重要农艺性状QTL分析
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摘要
大白菜（Brassica campestris L. ssp. pekenensis）起源于中国，是我国栽培面积最大的蔬菜作物，其很多农艺性状都是数量性状。构建高密度的永久分子遗传图谱，并对重要农艺数量性状进行QTL定位，对于大白菜基因组学研究有重要意义。本研究在优化大白菜游离小孢子培养技术体系的基础上，创建了包括192个DH系的大白菜永久性作图群体；采用SSR和SRAP标记相结合的方法，构建了大白菜分子遗传图谱，对重要农艺性状进行了定位，主要研究结果如下：
1.为创制大白菜DH系构建永久作图群体，优化了大白菜游离小孢子培养体系。以13个大白菜杂交种为试材，对游离小孢子培养的关键技术进行了系统研究。不同基因型材料间小孢子胚状体发生率不同，小孢子胚状体发生率最高的为10.6胚/蕾，一个基因型不能产生胚状体。高温热激处理对提高胚状体诱导率作用显著，33℃处理24h或48h对于胚状体发生比72h更为有利。在NLN-13培养基中加入6-BA和NAA可以显著提高胚状体发生率，以‘0.1mg·L~(-1)6-BA+0.05mg·L~(-1)NAA’效果最好，胚状体发生率是未添加激素的2~3倍。抗生长素PCIB对提高胚状体发生率和直接成苗率具有显著作用，最佳的浓度是40μM，可使胚状体发生率提高3.4~6.2倍，直接成苗率提高9.6倍，PCIB对胚状体形成次生胚也都作用。胚状体转胚时间以21d胚龄为宜。本研究成苗率最高的基因型可以达到67.4%，平均自然加倍率可以达到57.5%。
2.用不同生态型的两个大白菜自交系配制的杂交种进行游离小孢子培养，获得了192个DH系。为其为作图群体，采用SSR和SRAP两种标记技术，通过Joinmap3.0软件构建了一张永久性的遗传连锁图，包含233个标记，SSR标记43个、SRAP标记190个，长度达到1063.8cM，平均图距为4.6cM，标记间的最大间距为18.0cM，最短的为0cM。共10个连锁群，每个连锁群上的标记数在14~32个之间，长度在38.2~150.5cM范围内，利用SSR标记锚定可以与染色体对应。在233个多态性标记中，149个(63.9%)显著偏离孟德尔1:1遗传分离比率，各连锁群上偏分离标记比例不同。
3.以233个分子标记的大白菜分子遗传图谱为框架，在沈阳和大连两地对大白菜的形态性状包括株高、株幅、外叶长、外叶宽、叶形指数、叶柄长、叶柄宽、叶柄厚8个性状进行了QTL定位和遗传效应分析。采用基于混合线性模型的复合区间作图法的QTLnetwork2.0软件，在9个连锁群上定位了37个形态学相关的QTL，检测到4对上位性QTL，分布在A05，A06，A08，A09连锁群上。其中控制株高的QTL有4个，控制株幅的QTL有3个，控制外叶长的QTL有7个，控制外叶宽的QTL有6个，控制叶形指数的QTL有4个，控制叶柄长的QTL有2个，控制叶柄宽的QTL有5个，控制叶柄厚的QTL有6个。各QTL贡献率不同，在5.09%~31.37%之间，24个QTL的贡献率大于10.00%。贡献率最高的是控制外叶宽的QTL，在A10连锁群上。相关性状QTL的位置往往集中在连锁群上相同或相近的区域，控制株高和外叶长的2个QTL位于A08的相同位置（73.9cM），控制外叶长和叶柄厚的2个QTL均位于A05连锁群的33.4cM处，控制外叶长和叶柄长的2个QTL均位于A10连锁群的0.0cM处，控制外叶宽和叶柄厚的2个QTL位于A06连锁群上12.7cM处。外叶宽和叶柄宽在两地都检测到相同位置的QTL。
4.通过沈阳、大连两地的田间性状调查的数据分析，利用已构建好的分子遗传图谱，在混合线性模型下，对叶片颜色相关性状进行QTL分析，检测到共51个QTL定位于7个连锁群上，并且检测到有7对上位性QTL。将控制外叶色L的QTL有4个，控制外叶色a的QTL有4个，控制外叶色b的QTL有4个、控制叶柄色L的QTL，控制叶柄色a的QTL有4个，控制叶柄色b的QTL6个、控制球帮色L的QTL有6个、控制球帮色a的QTL有4个，控制球帮色b的QTL有5个，控制球心色L的QTL有5个，控制球心色a的QTL有3个，控制球心色b的QTL有3个。各QTL贡献率不同，在2.12%~73.19%之间,12个QTL的贡献率大于30.00%。在A03连锁群上0.0~4.0cM区域，检测到控制叶柄色L，叶柄色a，叶柄色b，球帮色L，球帮色a，球帮色b的QTL共12个，解释的遗传贡献率为34.54%~73.19%，且很多QTL都在两个地点检测到，此区域存在着控制叶柄色和球帮色重要的QTL。
Chinese cabbage (Brassica campestris L. ssp. pekenensis) is originated from China, which has thelargest cultivation area. Many important agronomic traits of Chinese cabbage are quantitative traits, so it isnecessary to construction a genetic map and map QTL on the linkages. The study tried to opitimize themicrospore culture technique and obtained a DH population. Based on the SSR and SRAP markers, agenetic map was constructed for mapping the important agronomic traits. Marker analysis was performedon192doubled haploid (DH) lines obtained from F1progeny of two homozygous parents. The mainresults of the present study were as follows:
1. In order to establish an efficient technique system of isolated microspore culture and constructionimmortal mapping population in Chinese cabbage, isolated microspore culture was performed using13F1hybrids of Chinese cabbage. Genotypic difference in embryogenic response was observed, the highestfrequency of embryogenesis was10.6embryos per bud, and one genotype did not response toembryogenesis. Heat shock treatment is efficient for embryogenesis, compared with the longer incubationperiod (72h), heat shock treatment at33℃for24or48h was more beneficial for microsporeembryogenesis. The embryogenesis frequency can improve2-3fold by adding ‘0.1mg·L~(-1)6-BA+0.05mg·L~(-1)NAA’to NLN-13medium. While anti-auxin PCIB is used to enhance microspore embryogenesisand plant regeneration without an intervening callus phase. The optimum concentration of PCIBapplication was found to be40μM in NLN-13medium, which resulted in a3.4-to6.2-fold increase in thenumber of embryos (8.27-19.2embryos per bud) and a9.6-fold increase (21.33%) in the plant regenerationfrequency in comparison with the controls. The embryos, produced in NLN medium supplemented with40μM PCIB and transferred at the21-day-old reached the high direct plant regeneration rate. The highestplant regeneration rate reached to67.4%and the average spontaneous doubling frequency of the differentgenotypes of B. rapa was57.5%.
2. A genetic linkage map of Chinese cabbage was constructed, which was based on a DH populationwith192inividuals. SSR and SRAP were adoptd and233markers including43SSR,190SRAP markerswere integrated into the resulted map using Joinmap3.0version. This map consisted of10linages groups,covering1063.8cM with a mean marker intervalof4.6cM, the largest interval between markers was18.0cM. Various linkage groups were feature by14-32markers,38.2-150.5cM length. The10linkagemaps can correspond to chromosomes by the anchor SSR markers. Among the233polymorphic markers,149(63.9%) deviated from the Mendelian segregation ratios.
3. Based on a linkage map with233markers, QTL network2.0is used to investigate QTL of themajor traits related to plant morphology using the data collected under two locations.37QTLs and4paisrof epistatic loci controlling8agronomic traits related to morphology traits were mapped on9linkages.These QTLs included4for plant heitht,3for plant diameter,7for leaf length,6for leaf width,4for leaflength/Leaf width,2for petiole length,5for petiole width,6for petiole thickness. The explained varianceswere different, ranged from5.09%-31.37%.24QTL can explain more than10.00%. The QTL controllingleaf width was mapped on A10. The QTL of assoeiated traits often located on the same loci or near regionof a linkage group.2QTLs controlling plant height and leaf length were mapped on the same positon ofA08(73.9cM),2QTLs controlling leaf length and petiole length were mapped on the same positon of A05(33.4cM),2QTLs controlling leaf length and petiole were mapped on the same positon of A10(0.0cM),2QTLs controlling leaf width and petiole thickness were mapped on the same positon of A06(12.7cM). TheQTLs controlling leaf width and petiole width were detected in the two locations.
4. Based on a linkage map with233markers, QTL network2.0is used to investigate QTL of the major traits related to leaf colour traits using the data collected under two locations.51QTLs and7paisr ofepistatic loci controlling colour traits were mapped on7linkages. These QTLs included4for outer leafcolour L,4for outer leaf colour a,4for outer leaf colour b,4for petiole colour1L,4for petiole colour1a,6for petiole colour1b,6for petiole colour2L,4for petiole colour2a,5for petiole colour2b,5for innerleaf colour L,3for inner leaf colour a,3for inner leaf colour b. The explained variances were different,ranged from2.12%~73.19%.12QTL can explain more than30.00%.12QTLs controlling petiole colourwere mapped between0.0~4.0cM on A03, each QTL can explained between34.54%~73.19%and manyQTLs were detected in the two locations, this area existed important QTL controlling petiole colour.
1.为创制大白菜DH系构建永久作图群体，优化了大白菜游离小孢子培养体系。以13个大白菜杂交种为试材，对游离小孢子培养的关键技术进行了系统研究。不同基因型材料间小孢子胚状体发生率不同，小孢子胚状体发生率最高的为10.6胚/蕾，一个基因型不能产生胚状体。高温热激处理对提高胚状体诱导率作用显著，33℃处理24h或48h对于胚状体发生比72h更为有利。在NLN-13培养基中加入6-BA和NAA可以显著提高胚状体发生率，以‘0.1mg·L~(-1)6-BA+0.05mg·L~(-1)NAA’效果最好，胚状体发生率是未添加激素的2~3倍。抗生长素PCIB对提高胚状体发生率和直接成苗率具有显著作用，最佳的浓度是40μM，可使胚状体发生率提高3.4~6.2倍，直接成苗率提高9.6倍，PCIB对胚状体形成次生胚也都作用。胚状体转胚时间以21d胚龄为宜。本研究成苗率最高的基因型可以达到67.4%，平均自然加倍率可以达到57.5%。
2.用不同生态型的两个大白菜自交系配制的杂交种进行游离小孢子培养，获得了192个DH系。为其为作图群体，采用SSR和SRAP两种标记技术，通过Joinmap3.0软件构建了一张永久性的遗传连锁图，包含233个标记，SSR标记43个、SRAP标记190个，长度达到1063.8cM，平均图距为4.6cM，标记间的最大间距为18.0cM，最短的为0cM。共10个连锁群，每个连锁群上的标记数在14~32个之间，长度在38.2~150.5cM范围内，利用SSR标记锚定可以与染色体对应。在233个多态性标记中，149个(63.9%)显著偏离孟德尔1:1遗传分离比率，各连锁群上偏分离标记比例不同。
3.以233个分子标记的大白菜分子遗传图谱为框架，在沈阳和大连两地对大白菜的形态性状包括株高、株幅、外叶长、外叶宽、叶形指数、叶柄长、叶柄宽、叶柄厚8个性状进行了QTL定位和遗传效应分析。采用基于混合线性模型的复合区间作图法的QTLnetwork2.0软件，在9个连锁群上定位了37个形态学相关的QTL，检测到4对上位性QTL，分布在A05，A06，A08，A09连锁群上。其中控制株高的QTL有4个，控制株幅的QTL有3个，控制外叶长的QTL有7个，控制外叶宽的QTL有6个，控制叶形指数的QTL有4个，控制叶柄长的QTL有2个，控制叶柄宽的QTL有5个，控制叶柄厚的QTL有6个。各QTL贡献率不同，在5.09%~31.37%之间，24个QTL的贡献率大于10.00%。贡献率最高的是控制外叶宽的QTL，在A10连锁群上。相关性状QTL的位置往往集中在连锁群上相同或相近的区域，控制株高和外叶长的2个QTL位于A08的相同位置（73.9cM），控制外叶长和叶柄厚的2个QTL均位于A05连锁群的33.4cM处，控制外叶长和叶柄长的2个QTL均位于A10连锁群的0.0cM处，控制外叶宽和叶柄厚的2个QTL位于A06连锁群上12.7cM处。外叶宽和叶柄宽在两地都检测到相同位置的QTL。
4.通过沈阳、大连两地的田间性状调查的数据分析，利用已构建好的分子遗传图谱，在混合线性模型下，对叶片颜色相关性状进行QTL分析，检测到共51个QTL定位于7个连锁群上，并且检测到有7对上位性QTL。将控制外叶色L的QTL有4个，控制外叶色a的QTL有4个，控制外叶色b的QTL有4个、控制叶柄色L的QTL，控制叶柄色a的QTL有4个，控制叶柄色b的QTL6个、控制球帮色L的QTL有6个、控制球帮色a的QTL有4个，控制球帮色b的QTL有5个，控制球心色L的QTL有5个，控制球心色a的QTL有3个，控制球心色b的QTL有3个。各QTL贡献率不同，在2.12%~73.19%之间,12个QTL的贡献率大于30.00%。在A03连锁群上0.0~4.0cM区域，检测到控制叶柄色L，叶柄色a，叶柄色b，球帮色L，球帮色a，球帮色b的QTL共12个，解释的遗传贡献率为34.54%~73.19%，且很多QTL都在两个地点检测到，此区域存在着控制叶柄色和球帮色重要的QTL。
Chinese cabbage (Brassica campestris L. ssp. pekenensis) is originated from China, which has thelargest cultivation area. Many important agronomic traits of Chinese cabbage are quantitative traits, so it isnecessary to construction a genetic map and map QTL on the linkages. The study tried to opitimize themicrospore culture technique and obtained a DH population. Based on the SSR and SRAP markers, agenetic map was constructed for mapping the important agronomic traits. Marker analysis was performedon192doubled haploid (DH) lines obtained from F1progeny of two homozygous parents. The mainresults of the present study were as follows:
1. In order to establish an efficient technique system of isolated microspore culture and constructionimmortal mapping population in Chinese cabbage, isolated microspore culture was performed using13F1hybrids of Chinese cabbage. Genotypic difference in embryogenic response was observed, the highestfrequency of embryogenesis was10.6embryos per bud, and one genotype did not response toembryogenesis. Heat shock treatment is efficient for embryogenesis, compared with the longer incubationperiod (72h), heat shock treatment at33℃for24or48h was more beneficial for microsporeembryogenesis. The embryogenesis frequency can improve2-3fold by adding ‘0.1mg·L~(-1)6-BA+0.05mg·L~(-1)NAA’to NLN-13medium. While anti-auxin PCIB is used to enhance microspore embryogenesisand plant regeneration without an intervening callus phase. The optimum concentration of PCIBapplication was found to be40μM in NLN-13medium, which resulted in a3.4-to6.2-fold increase in thenumber of embryos (8.27-19.2embryos per bud) and a9.6-fold increase (21.33%) in the plant regenerationfrequency in comparison with the controls. The embryos, produced in NLN medium supplemented with40μM PCIB and transferred at the21-day-old reached the high direct plant regeneration rate. The highestplant regeneration rate reached to67.4%and the average spontaneous doubling frequency of the differentgenotypes of B. rapa was57.5%.
2. A genetic linkage map of Chinese cabbage was constructed, which was based on a DH populationwith192inividuals. SSR and SRAP were adoptd and233markers including43SSR,190SRAP markerswere integrated into the resulted map using Joinmap3.0version. This map consisted of10linages groups,covering1063.8cM with a mean marker intervalof4.6cM, the largest interval between markers was18.0cM. Various linkage groups were feature by14-32markers,38.2-150.5cM length. The10linkagemaps can correspond to chromosomes by the anchor SSR markers. Among the233polymorphic markers,149(63.9%) deviated from the Mendelian segregation ratios.
3. Based on a linkage map with233markers, QTL network2.0is used to investigate QTL of themajor traits related to plant morphology using the data collected under two locations.37QTLs and4paisrof epistatic loci controlling8agronomic traits related to morphology traits were mapped on9linkages.These QTLs included4for plant heitht,3for plant diameter,7for leaf length,6for leaf width,4for leaflength/Leaf width,2for petiole length,5for petiole width,6for petiole thickness. The explained varianceswere different, ranged from5.09%-31.37%.24QTL can explain more than10.00%. The QTL controllingleaf width was mapped on A10. The QTL of assoeiated traits often located on the same loci or near regionof a linkage group.2QTLs controlling plant height and leaf length were mapped on the same positon ofA08(73.9cM),2QTLs controlling leaf length and petiole length were mapped on the same positon of A05(33.4cM),2QTLs controlling leaf length and petiole were mapped on the same positon of A10(0.0cM),2QTLs controlling leaf width and petiole thickness were mapped on the same positon of A06(12.7cM). TheQTLs controlling leaf width and petiole width were detected in the two locations.
4. Based on a linkage map with233markers, QTL network2.0is used to investigate QTL of the major traits related to leaf colour traits using the data collected under two locations.51QTLs and7paisr ofepistatic loci controlling colour traits were mapped on7linkages. These QTLs included4for outer leafcolour L,4for outer leaf colour a,4for outer leaf colour b,4for petiole colour1L,4for petiole colour1a,6for petiole colour1b,6for petiole colour2L,4for petiole colour2a,5for petiole colour2b,5for innerleaf colour L,3for inner leaf colour a,3for inner leaf colour b. The explained variances were different,ranged from2.12%~73.19%.12QTL can explain more than30.00%.12QTLs controlling petiole colourwere mapped between0.0~4.0cM on A03, each QTL can explained between34.54%~73.19%and manyQTLs were detected in the two locations, this area existed important QTL controlling petiole colour.
引文
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