水稻细菌性条斑病抗性标记辅助选择研究
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摘要
分子标记辅助选择(Marker-assisted Selection,MAS)应用在水稻质量性状的辅助选择已比较成熟,但对数量性状的辅助选择还在探索中。本研究对利用分子标记辅助选择改良由多基因控制的水稻细菌性条斑病抗性的有关问题进行了研究: 1)在筛选与抗病基因紧密连锁的SSR分子标记的基础上,通过回交或混合选择结合MAS技术的方法,把来自水稻品种Dular中的细条病抗性QTL qBlsr-11-1导入新的遗传背景中,并对分子标记辅助选择的效果作了评价;2)对利用标记辅助聚合技术选育抗细条病新品种的可行性进行分析,探讨在水稻细条病抗病育种过程中抗性鉴定和标记辅助选择如何有效结合的问题。主要结果如下:
1、在前人对水稻细条病抗性QTL qBlsr5a、qBlsr3d、qBlsr5b和qBlsr-11-1定位的基础上,利用已公布的水稻基因组信息,获得了2个与qBlsr5a紧密连锁的标记RM153和RM17769; 2个与qBlsr3d紧密连锁的分子标记RM523和RM231;2个与qBlsr5b紧密连锁的标记RM440和RM173。这6个标记均在抗性亲本Dular和H359R间存在多态性。得到3个与qBlsr-11-1紧密连锁的标记RM120、RM26267和RM552,其中RM120和RM26267在供体亲本Dular、H359R和受体亲本Ⅱ-32B间存在多态性,RM26267和RM552在供体亲本Dular和受体亲本明恢2155间存在多态性。上述标记与相应的目标QTL的推测距离在0.4-4.9cM之间,基本可以满足MAS对标记选择准确率的要求。
2、在利用回交结合MAS技术把qBlsr-11-1导入II-32B的过程中,运用位于该目标QTL双侧紧密连锁的标记RM120和RM26267,或仅用RM120对杂合抗病标记基因型进行选择均有效,但利用双标记选择的效果比单标记的选择效果好;而在该回交群体中仅用RM26267单独对杂合抗病标记基因型进行选择是无效的。
3、在Dular×明恢2155的F3:4群体中,运用双标记RM552和RM26267或各单标记选出的纯合抗病标记基因型(BBCC、CC和BB)间的病斑平均值差异不显著,但是它们与感病亲本明恢2155、三种杂合标记基因型(BbCc,Bb和Cc)及三种纯合感病标记基因型(cc,bb和bbcc)的病斑长度差异均达极显著水平。BbCc和Cc这两种杂合标记基因型与纯合感病标记基因型(cc,bb和bbcc)和感病亲本明恢2155的病斑长度差异也均达到极显著水平。但单独利用RM26267选出的杂合标记基因型与纯合感病基因型的病斑长度差异不显著。
4、先进行抗性鉴定再进行标记基因型选择或直接进行标记基因型选择两种不同的途径均可以获得携带4个、3个、2个抗性QTL的个体,但效率不同。仅在30株极端抗病个体中即可选出含有4个和3个抗性QTL的纯合抗病标记基因型单株3株和4株,而利用标记基因型选择从未抗性鉴定的300株的群体中,仅选出含4个和3个QTL抗性QTL的纯合抗病标记基因型单株1株和7株。可见,在利用MAS技术对数量抗病性状的改良中,在抗性鉴定结果较理想的情况下,先进行抗性鉴定再进行标记基因型选择,能选育出更多聚合多个抗性QTL的抗病单株,并且可以大大减小分子标记的工作量和费用。
5、含有2个以上抗性QTL的单株(株系),其抗性均比感病亲本H359有极显著提高。含有4个抗性QTL的L1类型和含有3个抗性QTL的L2、L3类型以及含有2个抗性QTL的L6类型内单株的总平均病斑长度与原始抗源Acc8558和Dular的抗性差异不显著,表明只要对已定位的2~4个细条病抗性QTL的聚合,即有可能选育出与抗源抗性相当的新品系。
Molecular marker-assisted selection (MAS) has been successfully used in improving qualitive traits of crops, and gradurally used in quantitative traits. In this study, we researched the genetic improvement of bacterial leaf streak disease in rice by means of molecular marker-assisted selection. 1) On the basis of screening SSR markers closely linked to resistant QTL, we introgressed the QTL qBlsr-11-1(from rice variety Dular)for the bacterial leaf streak resistance into new genetic backgrounds of rice by means of backcross method combined with MAS or by bulk method combined with MAS,and evaluated the efficiency of MAS. 2) We analyzied the feasibility to pyramid QTLs for improving bacterial leaf streak resistance by the use of marker-assisted selection, and explored some issues concerning breeding of bacterial leaf streak resistance. The main results are as follows:
1. On the basis of located QTL qBlsr5a, qBlsr3d, qBlsr5b and qBlsr-11-1 for bacterial leaf streak resistance and the rice genome information published before, we obtained two SSR markers RM153 and RM17769 closely linked to the qBlsr5a, two SSR markers RM523 and RM231 closely linked to the qBlsr3d, and two SSR markers RM440 and RM173 closely linked to the qBlsr5b. These six SSR markers perform polymorphisms between the donor parents Dular and H359R. We also got three SSR markers RM120, RM26267and RM552 linked to the qBlsr-11-1. The markers RM120 and RM26267 exist polymorphisms between two donor parents Dular and H359R and the recipient parentⅡ-32B; the markers RM26267 and RM552 exist polymorphisms between the donor parental Dular and the recipient parent Minghui 2155. Their distances from the markers to the target QTLs estimated at 0.4-4.9cM, which will meet the basic requirements for MAS.
2. In the prosess of introgressing qBlsr-11-1 into II-32B by combining backcross method with MAS, both of selection methods are efficiency by use either duoble side markers RM120 and RM26267or one side marker RM120 closely linked the qBlsr-11-1 to select the resistantly heterozygous marker genotypes; but selection is not efficiency by use the marker RM26267 only.
3. In the F3:4 population derived from Dular×Minghui 2155, the difference was not significant in average lesion lengths of homozygous resistant genotypes (BB, CC and BBCC) selected by use both double side markers RM552 and RM26267 or one side marker. But the average lesion lengths were significantly difference between the homozygous resistant genotypes (BBCC,CC and BB) and infectious parent Minghui 2155, the heterozygous marker genotypes (BbCc, Bb and Cc), and the susceptibly homozygous marker genotypes (cc, bb and bbcc). The average lesion lengths were significantly difference between heterozygous marker genotype (BbCc and Cc) and the susceptibly homozygous marker genotypes (cc, bb and bbcc) and the infectious parent Minghui 2155 too. The difference was not significant in average lesion lengths between heterozygous marker genotype selected by RM26267 marker and the infectious parent Minghui 2155.
4. We could obtain resistant plants with 4 resistant QTLs, 3 resistant QTLs and 2 resistant QTLs respectively, by means of the methods both through identifing disease resistance first and then select the marker genotype or direct select the marker genotype, but the efficiency is different. We obtained homozygous marker genotypes 3 plants and 4 plants which were with 4 resistant QTLs and 3 resistant QTLs respectively, in the 30 extremely resistant plants. And by the method of direct marker genotypes selection,we obtained homozygous marker genotype 1 plants and 7 plants with 4 resistant QTLs and 3 resistant QTLs respectively, in the population comprises 300 plants. Above results indicated that,it would be an ideal method to identify disease resistance first and then select the marker genotype, in improving the quantitative traits such as bacterial leaf streak resistance, as this could select more ideal plants and greatly reduce work and costs.
5. The plants(lines)containing more than 2 resisitant QTLs were significantly higher than susceptible parent H359 in resistant level. The difference was not significant in average lesion lengths among the plants carrying 4 resistant QTL (L1), 3 resistant QTL (L2 and L3), 2 resistant QTL (L6) and their resistant donor parents Acc8558 and Dular. The results indicating that, we can breed a new rice variety similar with their resistant donor parents in resistant level, as long as 2 ~ 4 resistant QTL been pyramided.
1、在前人对水稻细条病抗性QTL qBlsr5a、qBlsr3d、qBlsr5b和qBlsr-11-1定位的基础上,利用已公布的水稻基因组信息,获得了2个与qBlsr5a紧密连锁的标记RM153和RM17769; 2个与qBlsr3d紧密连锁的分子标记RM523和RM231;2个与qBlsr5b紧密连锁的标记RM440和RM173。这6个标记均在抗性亲本Dular和H359R间存在多态性。得到3个与qBlsr-11-1紧密连锁的标记RM120、RM26267和RM552,其中RM120和RM26267在供体亲本Dular、H359R和受体亲本Ⅱ-32B间存在多态性,RM26267和RM552在供体亲本Dular和受体亲本明恢2155间存在多态性。上述标记与相应的目标QTL的推测距离在0.4-4.9cM之间,基本可以满足MAS对标记选择准确率的要求。
2、在利用回交结合MAS技术把qBlsr-11-1导入II-32B的过程中,运用位于该目标QTL双侧紧密连锁的标记RM120和RM26267,或仅用RM120对杂合抗病标记基因型进行选择均有效,但利用双标记选择的效果比单标记的选择效果好;而在该回交群体中仅用RM26267单独对杂合抗病标记基因型进行选择是无效的。
3、在Dular×明恢2155的F3:4群体中,运用双标记RM552和RM26267或各单标记选出的纯合抗病标记基因型(BBCC、CC和BB)间的病斑平均值差异不显著,但是它们与感病亲本明恢2155、三种杂合标记基因型(BbCc,Bb和Cc)及三种纯合感病标记基因型(cc,bb和bbcc)的病斑长度差异均达极显著水平。BbCc和Cc这两种杂合标记基因型与纯合感病标记基因型(cc,bb和bbcc)和感病亲本明恢2155的病斑长度差异也均达到极显著水平。但单独利用RM26267选出的杂合标记基因型与纯合感病基因型的病斑长度差异不显著。
4、先进行抗性鉴定再进行标记基因型选择或直接进行标记基因型选择两种不同的途径均可以获得携带4个、3个、2个抗性QTL的个体,但效率不同。仅在30株极端抗病个体中即可选出含有4个和3个抗性QTL的纯合抗病标记基因型单株3株和4株,而利用标记基因型选择从未抗性鉴定的300株的群体中,仅选出含4个和3个QTL抗性QTL的纯合抗病标记基因型单株1株和7株。可见,在利用MAS技术对数量抗病性状的改良中,在抗性鉴定结果较理想的情况下,先进行抗性鉴定再进行标记基因型选择,能选育出更多聚合多个抗性QTL的抗病单株,并且可以大大减小分子标记的工作量和费用。
5、含有2个以上抗性QTL的单株(株系),其抗性均比感病亲本H359有极显著提高。含有4个抗性QTL的L1类型和含有3个抗性QTL的L2、L3类型以及含有2个抗性QTL的L6类型内单株的总平均病斑长度与原始抗源Acc8558和Dular的抗性差异不显著,表明只要对已定位的2~4个细条病抗性QTL的聚合,即有可能选育出与抗源抗性相当的新品系。
Molecular marker-assisted selection (MAS) has been successfully used in improving qualitive traits of crops, and gradurally used in quantitative traits. In this study, we researched the genetic improvement of bacterial leaf streak disease in rice by means of molecular marker-assisted selection. 1) On the basis of screening SSR markers closely linked to resistant QTL, we introgressed the QTL qBlsr-11-1(from rice variety Dular)for the bacterial leaf streak resistance into new genetic backgrounds of rice by means of backcross method combined with MAS or by bulk method combined with MAS,and evaluated the efficiency of MAS. 2) We analyzied the feasibility to pyramid QTLs for improving bacterial leaf streak resistance by the use of marker-assisted selection, and explored some issues concerning breeding of bacterial leaf streak resistance. The main results are as follows:
1. On the basis of located QTL qBlsr5a, qBlsr3d, qBlsr5b and qBlsr-11-1 for bacterial leaf streak resistance and the rice genome information published before, we obtained two SSR markers RM153 and RM17769 closely linked to the qBlsr5a, two SSR markers RM523 and RM231 closely linked to the qBlsr3d, and two SSR markers RM440 and RM173 closely linked to the qBlsr5b. These six SSR markers perform polymorphisms between the donor parents Dular and H359R. We also got three SSR markers RM120, RM26267and RM552 linked to the qBlsr-11-1. The markers RM120 and RM26267 exist polymorphisms between two donor parents Dular and H359R and the recipient parentⅡ-32B; the markers RM26267 and RM552 exist polymorphisms between the donor parental Dular and the recipient parent Minghui 2155. Their distances from the markers to the target QTLs estimated at 0.4-4.9cM, which will meet the basic requirements for MAS.
2. In the prosess of introgressing qBlsr-11-1 into II-32B by combining backcross method with MAS, both of selection methods are efficiency by use either duoble side markers RM120 and RM26267or one side marker RM120 closely linked the qBlsr-11-1 to select the resistantly heterozygous marker genotypes; but selection is not efficiency by use the marker RM26267 only.
3. In the F3:4 population derived from Dular×Minghui 2155, the difference was not significant in average lesion lengths of homozygous resistant genotypes (BB, CC and BBCC) selected by use both double side markers RM552 and RM26267 or one side marker. But the average lesion lengths were significantly difference between the homozygous resistant genotypes (BBCC,CC and BB) and infectious parent Minghui 2155, the heterozygous marker genotypes (BbCc, Bb and Cc), and the susceptibly homozygous marker genotypes (cc, bb and bbcc). The average lesion lengths were significantly difference between heterozygous marker genotype (BbCc and Cc) and the susceptibly homozygous marker genotypes (cc, bb and bbcc) and the infectious parent Minghui 2155 too. The difference was not significant in average lesion lengths between heterozygous marker genotype selected by RM26267 marker and the infectious parent Minghui 2155.
4. We could obtain resistant plants with 4 resistant QTLs, 3 resistant QTLs and 2 resistant QTLs respectively, by means of the methods both through identifing disease resistance first and then select the marker genotype or direct select the marker genotype, but the efficiency is different. We obtained homozygous marker genotypes 3 plants and 4 plants which were with 4 resistant QTLs and 3 resistant QTLs respectively, in the 30 extremely resistant plants. And by the method of direct marker genotypes selection,we obtained homozygous marker genotype 1 plants and 7 plants with 4 resistant QTLs and 3 resistant QTLs respectively, in the population comprises 300 plants. Above results indicated that,it would be an ideal method to identify disease resistance first and then select the marker genotype, in improving the quantitative traits such as bacterial leaf streak resistance, as this could select more ideal plants and greatly reduce work and costs.
5. The plants(lines)containing more than 2 resisitant QTLs were significantly higher than susceptible parent H359 in resistant level. The difference was not significant in average lesion lengths among the plants carrying 4 resistant QTL (L1), 3 resistant QTL (L2 and L3), 2 resistant QTL (L6) and their resistant donor parents Acc8558 and Dular. The results indicating that, we can breed a new rice variety similar with their resistant donor parents in resistant level, as long as 2 ~ 4 resistant QTL been pyramided.
引文
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