分子标记辅助选择改良蜀恢527对白叶枯病的抗性和萍乡核不育水稻育性相关基因的定位分析
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摘要
Ⅰ
稻白叶枯病是水稻三大病害之一，对我国的水稻生产危害十分严重。防病治病最经济有效的措施是培育和种植抗病品种，将多个抗性基因聚合于一个品种，是提高品种抗性和延缓品种抗性“丧失”的有效途径。我国杂交稻生产面积大，抗性差，改良其抗性最直接有效的方法之一是在恢复系中导入多个显牲广谱抗性基因。Xa21是一个显性广谱抗性基因，Xa4能抵御我国大部分菌系，与这两个抗性基因紧密连锁的分子标记已经获得，使得用分子标记聚合这两个基因成为可能。
蜀恢527(527R)是四川农业大学水稻所育成的一个强恢复系，不抗白叶枯病。本研究用IRBB60(含Xa21、Xa4、xa5、xa13)为供体，用分子标记将Xa21和Xa4聚于527R，改良其抗性，结果如下：
与Xa21和Xa4紧密连锁的标记pTA248和MP12在亲本527R和IRBB60间有多态性，pTA248在(527R／IRBB21)F_2和MP12在(527R／IRBB4)F_2群体中符合1：2：1的分离比，是共显性标记。pTA248对Xa21选择的准确率≥97.1％，MP12对Xa4选择的准确率≥183.3％，准确率较高，可以用这两个标记对Xa21和Xa4进行辅助选择。
以527R为轮回受体亲本，IRBB60为供体亲本，经过3次回交一次自交，用分子标记结合田间选择，在BC_3F_2代选得10株Xa21和Xa4位点纯合，田间表现与527R相似的单株。编号为Line1～ine10。用分别与xa13和xa5连锁的标记RM264和RM611进行PCR分析，表明10个单株均不含xa13和xa5。接种鉴定表明，527R对P1、P6和CⅠ～CⅦ的代表菌系均表现感病，而Line1～ine10均表现抗。用在亲本间表现多态性的SSR标记和100个RAPD引物进行遗传背景分析发现，Line3检测到一个SSR差异位点，Line4、5、8、9、10检测到2个SSR差异位点。RAPD带型的相似率在92.1％～97.5％之间，其中Line3、5、6、8、9、10的相同率较高，在96.0％以上。综合两种分子分析结果Line3、5、8、9和10与蜀恢527的遗传背景最相似。
在自然不发病的环境下考察10个改良系及其与D62A和G46A配制组合的播抽期、株高、有效穗、千粒重、每穗颖花数、每穗粒数、结实率和产量8个农艺形状。结果表明：改良系的结实率达0.05显著，与对照527R相比，只有Line7的结实率高于527R达0.05显著，其余株系和性状表现与蜀恢527无明显差异。改良系与D62A所配组合在考察的8个性状中，无显著差异；但与G46A所配组合的千粒重、每穗颖花数和产量达统计上的显著，Line1，Line2和Line8与G46A所配组合与(G46A／527R)相比在上述几个性状上有显著差异，其它株系所配组合和性状没有达到统计上的差异。根据遗传和农艺性状分析的结果，Line5、Line9和line10与蜀恢527的遗传背景最相似。
进一步考察Lines、Lineg和linelo与G46A和D62A配制组合的抗性和产量构成性状表现，结
果表明，所配组合对P6不[1 Cl一CVll的混合菌液均表现抗，而对照(D62^/527R)和(G46A/527R)
表现感:在不发病环境下，各组合与对照无明显差异，但在分别接种P6和Cl~C妞的混合菌液发
病环境下，改良系所配组合明显优于对照，具体如下:与G46A配制的组合接种p6发病环境下，
千粒重和单株产量高于对照G46A/527R，分别达0.05和0.01显著，接种混合菌液发病环境下，穗
粒数、千粒重和单株产量高于G46A/527R，达0.01极显著。与D62A所配组合在接种P6和混合菌
液发病的环境下，千粒重和单株产量均高于对照062A/527R，达0.01极显著。由此可以将Lines、
Lineg和linelo作为改良的抗白叶枯病蜀恢527，进行广泛的配合力测试。
通过上述抗性鉴定和农艺形状考察，说明用分子标记将方公21和Xa4聚于527R，改良对白叶枯
病的抗性是成功的。
I
Bacterial Blight (BB) caused by Xanthomonas oryzae pv. Oryzae (Xoo) is one of the most destructive diseases of rice throughout the world. The most effective and economical approach to combat BB is breeding and use resistant varieties and one way to delay breakdown of rice resistance caused by BB and increase BB resistance of varieties is to pyramid multiple resistance genes into one rice variety . Hybrid rice is widely used in China but most of them are susceptible to BB, one of the most effective method to improve their resistance BB is to introgress two or more dominant resistant BB genes. Xa21 is one of the broad spectrum resistance gene Xa4 can control most of BB race in China, the molecular markers tightly linked with the two genes are developed, which make it possible to pyramid the two genes by molecular marker- assisted selection (MAS).
Shuhui527 (527R) is a promising restorer line bred by Rice Research Institute of Sichuan Agricultural University in recent years. However, it is susceptible to Bacterial Blight (BB) ,which limits its use widely. The objective of this study is improvement its resistance to BB using IRBB60,carrying resistant genes, Xa21,Xa4,xal3 and xa5,as donor parent by MAS. The main results are following:
pTA248 and Mpl2,tightly linked with Xa21 and Xa4,respectively,have polymorphism between 527R and IRBB60, and the segregate ratio of pTA248 in population of (527R/IRBB21)F2,MP12 in (527R/IRBB4)F2 all fit to 1 : 2 : 1,which indicate they are co-dominant markers. The selection accuracies of pTA248 and Mp12 are 97.1% and 83.3%,respectively.
Using 527R as recurrent receiptor parent, IRBB60 as donor parent,through three times backcrosses and one time self-cross, 10 plants.Linel-Line 10, were obtained which their loci Xa21 and Xa4 are homozygous and field phenotype are similar to those of 527R,using the MAS together with field phenotype selection. The PCR analysis with RM264 and RM611, tightly linked with xa13 and xa5, respectively, shows that the 10 plants have not xa13 and xa5. The resistance analysis indicates that the 10 plants(lines) are resistance to race P1,P6 and C I ~CVII.The genetic background analysis with SSR markers which have polymorphism between 527R and IRBB60 and 100 RAPD markers shows that the Line3, Line5, Line8, Line9 and Line10 are the most similar to 527R, which demonstrate that the genetic background recover rapidly through MAS coupled with strict field phenotype selection in every generation.
Under no diseased condition in Wenjiang, the agronomic traits of 10 improved lines and 527R and their hybrid combinations crossing with CMS D62A and G46A ,were investigated, which include days to heading ,1000-grain weight, plant height, panicles per plant, spikelets per panicle, grain per panicle , setting percentage and yield. The results show that setting percentage is significant at 0.05 level in 527R and 10 improved lines. The variation of the traits observed the hybrid combinations that each improved lines cross with D62A are not significant, but the analysis of the hybrid combinations that improved lines cross with G46A show that 1000-grain weight is significant at 0.05 level and the spikletes per panicles and yield are significant at 0.01 level. From the analysis of above ,the Line5,Line9 and Line 10 are the most similar to 527R.
The Line5,Line9 and LinelO were more evaluated under heavily diseased conditions caused by artificial inoculation P6 and mixed races of C I ~CVII in Hai'nan. The results show that their combinations which the 3 lines crossed with susceptible CMS D62A and G46A resistance to P6 and mixed race of C I ~CVII .The agronomic traits ,1000-gain weight, and yield per plant of the above combinations,are better than those of controls 'Gangyou527' and 'Dyou527' .Then the Line5,Line9 and Line10 can be used as the improved 527R resistant to BB ,and the more test are on the way.
The above analysis indicate it is success to improve the resistance to BB of 527R by pyramiding Xa21 and Xa4 using MAS.
稻白叶枯病是水稻三大病害之一，对我国的水稻生产危害十分严重。防病治病最经济有效的措施是培育和种植抗病品种，将多个抗性基因聚合于一个品种，是提高品种抗性和延缓品种抗性“丧失”的有效途径。我国杂交稻生产面积大，抗性差，改良其抗性最直接有效的方法之一是在恢复系中导入多个显牲广谱抗性基因。Xa21是一个显性广谱抗性基因，Xa4能抵御我国大部分菌系，与这两个抗性基因紧密连锁的分子标记已经获得，使得用分子标记聚合这两个基因成为可能。
蜀恢527(527R)是四川农业大学水稻所育成的一个强恢复系，不抗白叶枯病。本研究用IRBB60(含Xa21、Xa4、xa5、xa13)为供体，用分子标记将Xa21和Xa4聚于527R，改良其抗性，结果如下：
与Xa21和Xa4紧密连锁的标记pTA248和MP12在亲本527R和IRBB60间有多态性，pTA248在(527R／IRBB21)F_2和MP12在(527R／IRBB4)F_2群体中符合1：2：1的分离比，是共显性标记。pTA248对Xa21选择的准确率≥97.1％，MP12对Xa4选择的准确率≥183.3％，准确率较高，可以用这两个标记对Xa21和Xa4进行辅助选择。
以527R为轮回受体亲本，IRBB60为供体亲本，经过3次回交一次自交，用分子标记结合田间选择，在BC_3F_2代选得10株Xa21和Xa4位点纯合，田间表现与527R相似的单株。编号为Line1～ine10。用分别与xa13和xa5连锁的标记RM264和RM611进行PCR分析，表明10个单株均不含xa13和xa5。接种鉴定表明，527R对P1、P6和CⅠ～CⅦ的代表菌系均表现感病，而Line1～ine10均表现抗。用在亲本间表现多态性的SSR标记和100个RAPD引物进行遗传背景分析发现，Line3检测到一个SSR差异位点，Line4、5、8、9、10检测到2个SSR差异位点。RAPD带型的相似率在92.1％～97.5％之间，其中Line3、5、6、8、9、10的相同率较高，在96.0％以上。综合两种分子分析结果Line3、5、8、9和10与蜀恢527的遗传背景最相似。
在自然不发病的环境下考察10个改良系及其与D62A和G46A配制组合的播抽期、株高、有效穗、千粒重、每穗颖花数、每穗粒数、结实率和产量8个农艺形状。结果表明：改良系的结实率达0.05显著，与对照527R相比，只有Line7的结实率高于527R达0.05显著，其余株系和性状表现与蜀恢527无明显差异。改良系与D62A所配组合在考察的8个性状中，无显著差异；但与G46A所配组合的千粒重、每穗颖花数和产量达统计上的显著，Line1，Line2和Line8与G46A所配组合与(G46A／527R)相比在上述几个性状上有显著差异，其它株系所配组合和性状没有达到统计上的差异。根据遗传和农艺性状分析的结果，Line5、Line9和line10与蜀恢527的遗传背景最相似。
进一步考察Lines、Lineg和linelo与G46A和D62A配制组合的抗性和产量构成性状表现，结
果表明，所配组合对P6不[1 Cl一CVll的混合菌液均表现抗，而对照(D62^/527R)和(G46A/527R)
表现感:在不发病环境下，各组合与对照无明显差异，但在分别接种P6和Cl~C妞的混合菌液发
病环境下，改良系所配组合明显优于对照，具体如下:与G46A配制的组合接种p6发病环境下，
千粒重和单株产量高于对照G46A/527R，分别达0.05和0.01显著，接种混合菌液发病环境下，穗
粒数、千粒重和单株产量高于G46A/527R，达0.01极显著。与D62A所配组合在接种P6和混合菌
液发病的环境下，千粒重和单株产量均高于对照062A/527R，达0.01极显著。由此可以将Lines、
Lineg和linelo作为改良的抗白叶枯病蜀恢527，进行广泛的配合力测试。
通过上述抗性鉴定和农艺形状考察，说明用分子标记将方公21和Xa4聚于527R，改良对白叶枯
病的抗性是成功的。
I
Bacterial Blight (BB) caused by Xanthomonas oryzae pv. Oryzae (Xoo) is one of the most destructive diseases of rice throughout the world. The most effective and economical approach to combat BB is breeding and use resistant varieties and one way to delay breakdown of rice resistance caused by BB and increase BB resistance of varieties is to pyramid multiple resistance genes into one rice variety . Hybrid rice is widely used in China but most of them are susceptible to BB, one of the most effective method to improve their resistance BB is to introgress two or more dominant resistant BB genes. Xa21 is one of the broad spectrum resistance gene Xa4 can control most of BB race in China, the molecular markers tightly linked with the two genes are developed, which make it possible to pyramid the two genes by molecular marker- assisted selection (MAS).
Shuhui527 (527R) is a promising restorer line bred by Rice Research Institute of Sichuan Agricultural University in recent years. However, it is susceptible to Bacterial Blight (BB) ,which limits its use widely. The objective of this study is improvement its resistance to BB using IRBB60,carrying resistant genes, Xa21,Xa4,xal3 and xa5,as donor parent by MAS. The main results are following:
pTA248 and Mpl2,tightly linked with Xa21 and Xa4,respectively,have polymorphism between 527R and IRBB60, and the segregate ratio of pTA248 in population of (527R/IRBB21)F2,MP12 in (527R/IRBB4)F2 all fit to 1 : 2 : 1,which indicate they are co-dominant markers. The selection accuracies of pTA248 and Mp12 are 97.1% and 83.3%,respectively.
Using 527R as recurrent receiptor parent, IRBB60 as donor parent,through three times backcrosses and one time self-cross, 10 plants.Linel-Line 10, were obtained which their loci Xa21 and Xa4 are homozygous and field phenotype are similar to those of 527R,using the MAS together with field phenotype selection. The PCR analysis with RM264 and RM611, tightly linked with xa13 and xa5, respectively, shows that the 10 plants have not xa13 and xa5. The resistance analysis indicates that the 10 plants(lines) are resistance to race P1,P6 and C I ~CVII.The genetic background analysis with SSR markers which have polymorphism between 527R and IRBB60 and 100 RAPD markers shows that the Line3, Line5, Line8, Line9 and Line10 are the most similar to 527R, which demonstrate that the genetic background recover rapidly through MAS coupled with strict field phenotype selection in every generation.
Under no diseased condition in Wenjiang, the agronomic traits of 10 improved lines and 527R and their hybrid combinations crossing with CMS D62A and G46A ,were investigated, which include days to heading ,1000-grain weight, plant height, panicles per plant, spikelets per panicle, grain per panicle , setting percentage and yield. The results show that setting percentage is significant at 0.05 level in 527R and 10 improved lines. The variation of the traits observed the hybrid combinations that each improved lines cross with D62A are not significant, but the analysis of the hybrid combinations that improved lines cross with G46A show that 1000-grain weight is significant at 0.05 level and the spikletes per panicles and yield are significant at 0.01 level. From the analysis of above ,the Line5,Line9 and Line 10 are the most similar to 527R.
The Line5,Line9 and LinelO were more evaluated under heavily diseased conditions caused by artificial inoculation P6 and mixed races of C I ~CVII in Hai'nan. The results show that their combinations which the 3 lines crossed with susceptible CMS D62A and G46A resistance to P6 and mixed race of C I ~CVII .The agronomic traits ,1000-gain weight, and yield per plant of the above combinations,are better than those of controls 'Gangyou527' and 'Dyou527' .Then the Line5,Line9 and Line10 can be used as the improved 527R resistant to BB ,and the more test are on the way.
The above analysis indicate it is success to improve the resistance to BB of 527R by pyramiding Xa21 and Xa4 using MAS.
引文
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