亚洲地区国内外大豆育成品种苗期耐淹性鉴定、种质优选和QTL关联定位
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摘要
洪涝灾害在世界范围内频繁发生,是世界上许多地区普遍发生的自然灾害,也是我国大豆生产主要的非生物逆境灾害之一。涝害情况分为湿/渍害和淹水,淹水也可分为全淹和半淹。研究表明大豆[Glycine max(L.)Merr.]苗期和花荚期对淹水反应较敏感,大豆不同品种之间耐淹性有差异,利用耐淹大豆品种是提高抗灾能力的有效途径,故发掘、鉴定耐淹资源及其有利基因服务大豆育种和生产实践具有重要的意义。
长江中下游地区常有梅雨天气,大豆苗期易形成涝害。在前人研究我国大豆资源耐淹性的基础上,本文进一步研究我国和亚洲其他国家大豆育成品种苗期耐淹性的遗传变异,发掘优异资源;对耐淹性进行关联定位,寻找优异等位变异及其载体材料,并追索在后代育成品种中的累积。同时,还试探研究半淹胁迫下的遗传机制,定位QTL,筛选与之紧密连锁的分子标记。所获主要结果如下:
1.选取来源于我国黄淮、中国南方、东亚(韩国和朝鲜)、东南亚(菲律宾、越南、泰国、印尼等)和南亚(印度、尼泊尔、巴基斯坦等)的亚洲大豆育成品种350份。采用Vo期全淹处理,相对死苗率为指标,以王芳等所定的一组对照种为参照,分析5个地区大豆育成品种耐淹性的遗传变异。结果表明:相对失绿率和相对萎蔫率与相对死苗率存在极显著相关(P＜.0001),可作为耐淹性的参考指标;以相对死苗率鉴定大豆品种的苗期耐淹性,亚洲大豆育成品种的耐淹性(109.8%)与地方品种群体的耐淹性(110.3%)相当,并均高于野生大豆群体(104.5%);中国黄淮、南方地区育成品种耐淹性变异大,变幅为4.8%-212.0%,高于亚洲引种材料群体,遗传变异丰富,覆盖了整个供试群体;各区均存在耐淹品种,优选出莒选23、南农493-1、黔豆2号等15份耐淹品种。优选品种与野生和地方种质相互补充,为大豆抗涝育种提供丰富的遗传种质。
2.对全部供试材料SSR标记的基因型数据进行遗传结构分析的结果,整个群体由两个差异明显的群体构成,这两个群体分别和国内材料及亚洲国外材料相对应,说明大豆从我国引入亚洲其它国家后育种方向的差异导致了群体遗传结构的根本性差异。因此需要分开研究。
3.对我国黄淮和南方187份育成品种(群体1)SSR标记基因型数据进行成对位点连锁不平衡、群体结构和耐淹性3个性状与标记的关联分析,结果表明:(1)该群体按遗传结构分为7个亚群,群体存在LD(Linkage Disequilibrium),不平衡成对位点所占比例较大(48.63%),LD衰减速率较快.(2)检测到与3个耐淹性状关联位点14个(Satt681、Satt373、Satt659等),同一位点与2个或3个性状关联,表明3个耐淹性状之间确有相关性.与前人结果比较无相同位点。(3)分别筛选出耐淹3个性状OTL的6个、10个和8个优异等位变异,优选出合豆2号、黔豆3号、诱变31和南农493-1等具有较多优异等位变异的载体材料。因此可通过聚合大豆优异基因提高耐淹性。
4.对亚洲地区国外104份材料(群体2)的耐淹性状与SSR标记关联分析发现:(1)与3个耐淹性状关联位点6个,(Sct033、Satt186、Satt269等)累计14个;存在同一位点与2个或者3个性状有关联。部分关联位点与群体定位结果一致。(2)分别筛选出耐淹3个性状QTL的8个、6个和8个优异等位变异,优选出PI208432、PI377576和PI481690等具有2个以上优异等位变异的耐淹典型载体材料。
同时还对群体2的百粒重、蛋白质和脂肪含量进行了与SSR标记的关联分析,与百粒重、蛋白质和脂肪关联的位点有21个,点累计29个。筛选出百粒重、蛋白质和脂肪性状QTL优异等位变异6个、8个和13个,获得PI175187、PI165926和PI210351A等优异载体材料。
5.本研究同时探索了半淹胁迫下的耐淹性遗传,选用“苏88-M21(较耐)×新沂小黑豆(高度敏感)”衍生的176个重组自交系(NJRISX)为材料,采取V2期保持土壤表层5-7cm水层淹水处理20天研究大豆耐淹性遗传规律。结果表明:(1)对8个可能和耐淹性有关性状中,确定相关性较高的3个(株高变量、叶龄和熟期株高)耐淹指数的均值评价家系耐淹性。(2)主基因+多基因混合遗传模型分析结果表明耐淹性受2对具有重叠作用的连锁主基因控制,并存在多基因互作。主基因遗传率为62.83%.(3)利用国家大豆改良中心提供的遗传图谱,采用Win QTL Cartographer 2.5开展了大豆耐淹的QTL定位研究.结果表明耐淹性存在超亲变异,家系间差异显著.利用CIM和MIM法共同检测到2个稳定位点,均位于连锁群L2上satt229-satt527和satt527-sat.286区间内,对表型变异的解释率为10.1%-25.2%。利用MIM方法还检测到1个QTL,也位于L2连锁群上。鉴于时间限制,全淹和半淹胁迫下大豆耐淹性的遗传差异及其关系有待进一步研究。
Many flooding disasters occur frequently due to the heavy rainfall and poor drainage encountered in many areas in the world, and flooding is a common production problem for soybeans [Glycine max (L.) Merr.] in China. Flooding has been recognized in two categories: waterlogging and saturation in soil, waterlogging ulteriorly recognized in small two categories: submergence and incomplete immergence. Soybean is sensitive to flooding and waterlogging stress, especially at early seedling stage as well as flowering and podding stages; On the other hand, there existed great difference in the tolerance to flooding or waterlogging in soybean germplasm, so utilizing tolerant cultivars is one of the most effective ways to improve the soybean production under flooding and waterlogging stresses condition. Identification and discovery of the tolerant germplasm and their target genes is important and basic research work for breeding purposes.
Soybean seedling stage is threatened by plum rains in the lower reaches of the Yangzi region. Based on the research of submergence tolerance in soybean, the main objectives of the present study were: (1) to evaluate and identify the submergence tolerance of released cultivar populations from different Asian regions at seedling stage to reveal their genetic variation; (2) to discover the tolerant gene loci and elite alleles to submergence as well as their carriers, and to reveal the cumulative characteristics of elite alleles in the five major pedigrees of Chinese cultivars through association mapping; (3)At the same time, to establish new method system and detect QTLs and linked molecular markers for incomplete immergence tolerance by using RIL population NJRLSX at V2 stage. The main results were as follows.
Total 350 accessions sampled from different eco-regions (Huang-Huai Valleys in China, Southern China, East Asia, southeast Asia, and south Asia) were tested for the genetic variation of submergence tolerance. A randomized block design experiment submerged at V0 stage of soybean seedling was carried by using relative death seedling rate(De), relative lost chlorophyll rate(Lo) and relative wilting rate(Wi) as indices and the germplasm which Wangfang has been ascertained as controls . The results included: (1) Lo and Wi correlated significantly with De (P<.0001) , and the indicator system involving De as major index and Lo, Wi as subsidiary indices was effective on the evaluation of submergence tolerance at seedling stage. The De value of Asia released cultivars were 109.8%, higher than that of wild soybeans (104.5%), and nearly equal to that of landrace (110.3%). Cultivars from China existed the highest variation among the different Asian regions, and the De range of Chinese cultivars covered the whole range of the sample population with the value from 4.8% to 212.0%. There existed tolerant variation in all tested regions, and 15 varieties with high tolerant to submergence were identified, i.e. Juxuan23, Nannong493-1, Qiandou2 Hao and so on. Among them, 7 from Huang-Huai, 4 from Southern China, 2 from south Asia, and 1 from East Asia and southeast Asia respectively. The selected tolerant cultivars can be new sources for tolerant breeding, and be compensatory with the tolerant accessions selected from landraces and annual wild soybean.
According to the analyses of genetic structure of the total of soybean accessions with 64 simple-sequence repeat (SSR) markers, these experimental materials can be divided into Chinese varieties (Population 1) and exotic varieties (Population 2). This indicated the difference of introduced soybeans breeding direction led to ultimate population genetic structure diversity, so need to study respectively.
The Population 1 contained 187 varieties from Huang-Huai valleys and southern China, and was scanned with 85 SSR markers distributed evenly the whole genome. The linkage disequilibrium (LD), population structure and association mapping of QTLs for submergence tolerance as well as the inheritance of the elite alleles in the main pedigree of Population lwere carried out by using Structure 2.2 and TASSEL software. It showed that (1) Population 1 was composed of seven subpopulations, LD was detected extensively (48.63%), and LD decays decreased quickly. (2) Total 14 SSR loci were detected to be associated with three submergence tolerance traits, and some loci associated simultaneously with two or three traits, i.e. Satt681、Satt373、Satt659 and so on, indicating the correlation among these traits. (3) Total 6, 10 and 8 Elite alleles and relative typical carriers of relative death seedling rate and other two traits (Lo and Wi) were also selected. Hedou2 Hao, Qiandou3 Hao, Youbian31 and Nannong493-1 possess more elite alleles. In the five pedigree populations, some elite alleles lost along with breeding cycle change, but others added, which imply the potentials of pyramiding different elite genes to improve the tolerance.
Association analysis of 64 SSR markers with the phenotypic data of 3 submergence tolerance traits, 100-seed weight, protein content and fat content was conducted in Population 2 which had 104 accessions from other Asian countries. the results were: (1) Six SSR loci (i.e. Sct033、Satt186、Satt269 and so on) with a total 14 loci were found to be associated with 3 submergence tolerance traits. There also found a certain number of loci associated simultaneously with two or more relative traits. Some loci were consistent with mapped QTLs from family-based linkage mapping procedure. (2) Total 8, 6 and 8 elite alleles of QTLs for submergence tolerance were identified respectively, and three accessions, i.e. PI208432、PI377576, PI481690, with 2 or more than 2 elite alleles were selected.
And the 21 SSR loci with a total 29 loci were found to be associated with 100-seed weight, protein content and fat content, some. For the traits of 100-seed weight, protein content and fat content, 6, 8 and 13 elite alleles were identified respectively, and three accessions, i.e. PI175187, PI165926 and PI210351A were selected.
RIL population NJRISX from Su88-M21 (tolerant)×Xinyixiaoheidou (Highly sensitive) was evaluated to reveal the genetic base of waterlogging tolerance. The waterlogging stress was kept 5-7 cm water above soil surface for 20 days at V2 stage. It showed: (1) Among the eight traits, the values of plant height increment, the age of leaves and plant height of mature appeared relatively stable and highly correlative, so the mean of values of these traits was used as the indicator to evaluate waterlogging tolerance. (2)The results from segregation analysis under the major gene plus polygene mixed inheritance model showed that waterlogging tolerance of soybean was controlled by two linkage major genes with duplicate effect and polygene detected, the heritability of major genes was 62.83%. (3) QTL mapping for waterlogging tolerance in soybean was carried out under the methods of composite interval mapping (CIM) and multiple interval mapping (MIM) of software Win QTL Cartographer 2.5. There existed transgressive segregation and significant different of lines in the NJRISX population. Two QTLs associated with waterlogging tolerance were identified through both CIM and MIM methods, and both located on linkage group L2 within the regions of Satt229～Satt527 and Satt527～satt286, respectively, explaining 10.1%～25.2% of the total phenotypic variation. With MIM method the other locus on linkage groups L2 was detected which need to be further verified.
长江中下游地区常有梅雨天气,大豆苗期易形成涝害。在前人研究我国大豆资源耐淹性的基础上,本文进一步研究我国和亚洲其他国家大豆育成品种苗期耐淹性的遗传变异,发掘优异资源;对耐淹性进行关联定位,寻找优异等位变异及其载体材料,并追索在后代育成品种中的累积。同时,还试探研究半淹胁迫下的遗传机制,定位QTL,筛选与之紧密连锁的分子标记。所获主要结果如下:
1.选取来源于我国黄淮、中国南方、东亚(韩国和朝鲜)、东南亚(菲律宾、越南、泰国、印尼等)和南亚(印度、尼泊尔、巴基斯坦等)的亚洲大豆育成品种350份。采用Vo期全淹处理,相对死苗率为指标,以王芳等所定的一组对照种为参照,分析5个地区大豆育成品种耐淹性的遗传变异。结果表明:相对失绿率和相对萎蔫率与相对死苗率存在极显著相关(P＜.0001),可作为耐淹性的参考指标;以相对死苗率鉴定大豆品种的苗期耐淹性,亚洲大豆育成品种的耐淹性(109.8%)与地方品种群体的耐淹性(110.3%)相当,并均高于野生大豆群体(104.5%);中国黄淮、南方地区育成品种耐淹性变异大,变幅为4.8%-212.0%,高于亚洲引种材料群体,遗传变异丰富,覆盖了整个供试群体;各区均存在耐淹品种,优选出莒选23、南农493-1、黔豆2号等15份耐淹品种。优选品种与野生和地方种质相互补充,为大豆抗涝育种提供丰富的遗传种质。
2.对全部供试材料SSR标记的基因型数据进行遗传结构分析的结果,整个群体由两个差异明显的群体构成,这两个群体分别和国内材料及亚洲国外材料相对应,说明大豆从我国引入亚洲其它国家后育种方向的差异导致了群体遗传结构的根本性差异。因此需要分开研究。
3.对我国黄淮和南方187份育成品种(群体1)SSR标记基因型数据进行成对位点连锁不平衡、群体结构和耐淹性3个性状与标记的关联分析,结果表明:(1)该群体按遗传结构分为7个亚群,群体存在LD(Linkage Disequilibrium),不平衡成对位点所占比例较大(48.63%),LD衰减速率较快.(2)检测到与3个耐淹性状关联位点14个(Satt681、Satt373、Satt659等),同一位点与2个或3个性状关联,表明3个耐淹性状之间确有相关性.与前人结果比较无相同位点。(3)分别筛选出耐淹3个性状OTL的6个、10个和8个优异等位变异,优选出合豆2号、黔豆3号、诱变31和南农493-1等具有较多优异等位变异的载体材料。因此可通过聚合大豆优异基因提高耐淹性。
4.对亚洲地区国外104份材料(群体2)的耐淹性状与SSR标记关联分析发现:(1)与3个耐淹性状关联位点6个,(Sct033、Satt186、Satt269等)累计14个;存在同一位点与2个或者3个性状有关联。部分关联位点与群体定位结果一致。(2)分别筛选出耐淹3个性状QTL的8个、6个和8个优异等位变异,优选出PI208432、PI377576和PI481690等具有2个以上优异等位变异的耐淹典型载体材料。
同时还对群体2的百粒重、蛋白质和脂肪含量进行了与SSR标记的关联分析,与百粒重、蛋白质和脂肪关联的位点有21个,点累计29个。筛选出百粒重、蛋白质和脂肪性状QTL优异等位变异6个、8个和13个,获得PI175187、PI165926和PI210351A等优异载体材料。
5.本研究同时探索了半淹胁迫下的耐淹性遗传,选用“苏88-M21(较耐)×新沂小黑豆(高度敏感)”衍生的176个重组自交系(NJRISX)为材料,采取V2期保持土壤表层5-7cm水层淹水处理20天研究大豆耐淹性遗传规律。结果表明:(1)对8个可能和耐淹性有关性状中,确定相关性较高的3个(株高变量、叶龄和熟期株高)耐淹指数的均值评价家系耐淹性。(2)主基因+多基因混合遗传模型分析结果表明耐淹性受2对具有重叠作用的连锁主基因控制,并存在多基因互作。主基因遗传率为62.83%.(3)利用国家大豆改良中心提供的遗传图谱,采用Win QTL Cartographer 2.5开展了大豆耐淹的QTL定位研究.结果表明耐淹性存在超亲变异,家系间差异显著.利用CIM和MIM法共同检测到2个稳定位点,均位于连锁群L2上satt229-satt527和satt527-sat.286区间内,对表型变异的解释率为10.1%-25.2%。利用MIM方法还检测到1个QTL,也位于L2连锁群上。鉴于时间限制,全淹和半淹胁迫下大豆耐淹性的遗传差异及其关系有待进一步研究。
Many flooding disasters occur frequently due to the heavy rainfall and poor drainage encountered in many areas in the world, and flooding is a common production problem for soybeans [Glycine max (L.) Merr.] in China. Flooding has been recognized in two categories: waterlogging and saturation in soil, waterlogging ulteriorly recognized in small two categories: submergence and incomplete immergence. Soybean is sensitive to flooding and waterlogging stress, especially at early seedling stage as well as flowering and podding stages; On the other hand, there existed great difference in the tolerance to flooding or waterlogging in soybean germplasm, so utilizing tolerant cultivars is one of the most effective ways to improve the soybean production under flooding and waterlogging stresses condition. Identification and discovery of the tolerant germplasm and their target genes is important and basic research work for breeding purposes.
Soybean seedling stage is threatened by plum rains in the lower reaches of the Yangzi region. Based on the research of submergence tolerance in soybean, the main objectives of the present study were: (1) to evaluate and identify the submergence tolerance of released cultivar populations from different Asian regions at seedling stage to reveal their genetic variation; (2) to discover the tolerant gene loci and elite alleles to submergence as well as their carriers, and to reveal the cumulative characteristics of elite alleles in the five major pedigrees of Chinese cultivars through association mapping; (3)At the same time, to establish new method system and detect QTLs and linked molecular markers for incomplete immergence tolerance by using RIL population NJRLSX at V2 stage. The main results were as follows.
Total 350 accessions sampled from different eco-regions (Huang-Huai Valleys in China, Southern China, East Asia, southeast Asia, and south Asia) were tested for the genetic variation of submergence tolerance. A randomized block design experiment submerged at V0 stage of soybean seedling was carried by using relative death seedling rate(De), relative lost chlorophyll rate(Lo) and relative wilting rate(Wi) as indices and the germplasm which Wangfang has been ascertained as controls . The results included: (1) Lo and Wi correlated significantly with De (P<.0001) , and the indicator system involving De as major index and Lo, Wi as subsidiary indices was effective on the evaluation of submergence tolerance at seedling stage. The De value of Asia released cultivars were 109.8%, higher than that of wild soybeans (104.5%), and nearly equal to that of landrace (110.3%). Cultivars from China existed the highest variation among the different Asian regions, and the De range of Chinese cultivars covered the whole range of the sample population with the value from 4.8% to 212.0%. There existed tolerant variation in all tested regions, and 15 varieties with high tolerant to submergence were identified, i.e. Juxuan23, Nannong493-1, Qiandou2 Hao and so on. Among them, 7 from Huang-Huai, 4 from Southern China, 2 from south Asia, and 1 from East Asia and southeast Asia respectively. The selected tolerant cultivars can be new sources for tolerant breeding, and be compensatory with the tolerant accessions selected from landraces and annual wild soybean.
According to the analyses of genetic structure of the total of soybean accessions with 64 simple-sequence repeat (SSR) markers, these experimental materials can be divided into Chinese varieties (Population 1) and exotic varieties (Population 2). This indicated the difference of introduced soybeans breeding direction led to ultimate population genetic structure diversity, so need to study respectively.
The Population 1 contained 187 varieties from Huang-Huai valleys and southern China, and was scanned with 85 SSR markers distributed evenly the whole genome. The linkage disequilibrium (LD), population structure and association mapping of QTLs for submergence tolerance as well as the inheritance of the elite alleles in the main pedigree of Population lwere carried out by using Structure 2.2 and TASSEL software. It showed that (1) Population 1 was composed of seven subpopulations, LD was detected extensively (48.63%), and LD decays decreased quickly. (2) Total 14 SSR loci were detected to be associated with three submergence tolerance traits, and some loci associated simultaneously with two or three traits, i.e. Satt681、Satt373、Satt659 and so on, indicating the correlation among these traits. (3) Total 6, 10 and 8 Elite alleles and relative typical carriers of relative death seedling rate and other two traits (Lo and Wi) were also selected. Hedou2 Hao, Qiandou3 Hao, Youbian31 and Nannong493-1 possess more elite alleles. In the five pedigree populations, some elite alleles lost along with breeding cycle change, but others added, which imply the potentials of pyramiding different elite genes to improve the tolerance.
Association analysis of 64 SSR markers with the phenotypic data of 3 submergence tolerance traits, 100-seed weight, protein content and fat content was conducted in Population 2 which had 104 accessions from other Asian countries. the results were: (1) Six SSR loci (i.e. Sct033、Satt186、Satt269 and so on) with a total 14 loci were found to be associated with 3 submergence tolerance traits. There also found a certain number of loci associated simultaneously with two or more relative traits. Some loci were consistent with mapped QTLs from family-based linkage mapping procedure. (2) Total 8, 6 and 8 elite alleles of QTLs for submergence tolerance were identified respectively, and three accessions, i.e. PI208432、PI377576, PI481690, with 2 or more than 2 elite alleles were selected.
And the 21 SSR loci with a total 29 loci were found to be associated with 100-seed weight, protein content and fat content, some. For the traits of 100-seed weight, protein content and fat content, 6, 8 and 13 elite alleles were identified respectively, and three accessions, i.e. PI175187, PI165926 and PI210351A were selected.
RIL population NJRISX from Su88-M21 (tolerant)×Xinyixiaoheidou (Highly sensitive) was evaluated to reveal the genetic base of waterlogging tolerance. The waterlogging stress was kept 5-7 cm water above soil surface for 20 days at V2 stage. It showed: (1) Among the eight traits, the values of plant height increment, the age of leaves and plant height of mature appeared relatively stable and highly correlative, so the mean of values of these traits was used as the indicator to evaluate waterlogging tolerance. (2)The results from segregation analysis under the major gene plus polygene mixed inheritance model showed that waterlogging tolerance of soybean was controlled by two linkage major genes with duplicate effect and polygene detected, the heritability of major genes was 62.83%. (3) QTL mapping for waterlogging tolerance in soybean was carried out under the methods of composite interval mapping (CIM) and multiple interval mapping (MIM) of software Win QTL Cartographer 2.5. There existed transgressive segregation and significant different of lines in the NJRISX population. Two QTLs associated with waterlogging tolerance were identified through both CIM and MIM methods, and both located on linkage group L2 within the regions of Satt229～Satt527 and Satt527～satt286, respectively, explaining 10.1%～25.2% of the total phenotypic variation. With MIM method the other locus on linkage groups L2 was detected which need to be further verified.
引文
1. Adler F R, Chase K, Cregan P B, et al. Genetics of soybean agronomic traits: I. comparison of three related recombinant inbred populations[J]. Crop Sci, 1999, 39: (6) 1642-1651.
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