九个重要小麦抗条锈病品种（系）抗病基因的遗传分析与分子作图
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摘要
小麦条锈病是由条锈菌(Puccinia striiformis f. sp. tritici)引起的世界性小麦病害,在我国曾多次大流行,给小麦生产造成重大损失。种植抗病品种是防治小麦条锈病最有效、经济和安全的措施。小麦对条锈菌的抗病性可以分为全生育期抗病性和成株期抗病性。全生育期抗病性又称苗期抗病性,这种抗病性通常是小种专化的,容易被新的毒性小种所克服,而成株期抗病性是由成株期表达的抗病基因所控制的,这种抗病性相对持久。高温抗条锈性是由较高的环境温度所诱导表达的一种非小种专化性的持久抗病性,分为全生育期和高温成株期抗病性两种类型。将各种抗病基因聚合,培育小麦抗病品种,将会对持久控制小麦条锈病发挥更大作用。因此,寻找和发掘新的小麦条锈病抗源、改良小麦品种抗锈性和促进小麦品种抗病基因的多样化,对控制小麦条锈病的流行和保障我国粮食生产安全具有重要意义。
本研究对普通小麦-华山新麦草易位系H9014-14-4-6-1、H9015-17-1-9-6、H9014-121-5-5-9、中梁12、中梁16、中梁21、三属麦1号和兰天1号抗条锈病基因进行了遗传分析以及分子作图,取得了以下结果：
1.对普通小麦-华山新麦草易位系H9014-14-4-6-1与铭贤169杂交的F1、F2和F3代在苗期温室条件下进行遗传分析,结果表明H9014-14-4-6-1对Sul1-4的抗病性由一对显性基因控制,暂命名为YrH9014；对CYR33的抗病性由一对隐形基因控制。利用群体分离分析法(BSA)筛选到与抗病基因YrH9014连锁的简单重复序列(SSR)标记。用中国春缺四体进一步确定了SSR标记所在染色体的具体位置,抗锈基因YrH9014被定位在2B短臂上。对F2分离群体的142个单株进行分析,构建了与YrH9014连锁的遗传图谱。系谱分析结合分子标记结果表明,YrH9014可能是来自于华山新麦草,并与已知抗条锈病基因不同的。
2.通过对H9015-17-1-9-6与铭贤169杂交的F1、F2和F3代进行遗传分析,表明H9015-17-1-9-6对CYR32的抗病性由一对显性基因控制,暂命名为YrH9015。利用BSA法对RGAP引物和SSR引物进行筛选,获得了与抗条锈病基因YrH9015连锁的7对RGAP标记和3对SSR标记,并将YrH9015定位于小麦5DL染色体。YrH9015可能来自于华山新麦草,并与已知抗条锈病基因不同。
3.对H9014-121-5-5-9与铭贤169杂交的F1、F2和F3代进行遗传分析,结果表明H9014-121-5-5-9对CYR31的抗病性由一对显性基因控制,暂命名为YrHA。利用BSA法对SSR引物进行筛选,获得了与YrHA连锁的7对SSR标记,并将YrHA定位于小麦1AL染色体。系谱分析结合分子标记结果表明,YrHA可能来自于华山新麦草,并与已知抗条锈病基因不同。
4.经典遗传学分析表明,中梁12对CYR30和CYR31抗病性分别由一对显性基因控制,分别暂命名为YrZhong12-1和YrZhong12-2。BSA法结合SSR标记分析,7AL染色体上的Xwmc695、Xcfd20、Xbarc121、Xbarc49与YrZhong12-1连锁；1AL染色体上的Xpsp3003、Xcfd2129、Xwmc673、Xwmc51与YrZhong12-2连锁。
5.采用生理小种CYR29和CYR30对中梁16进行遗传分析,结果表明中梁16对CYR29和CYR30抗病性分别由一对显性基因控制,分别暂命名为YrZhong16-1和YrZhong16-2。利用BSA法对SSR引物进行筛选,获得了5个SSR标记,并将YrZhong16-1定位在2AS染色体上；5个SSR标记与YrZhong16-2连锁,并将YrZhong16-2定位在7BL染色体上。
6.采用CYR30小种对中梁21进行遗传分析,并利用SSR分子标记进行遗传作图,发现中梁21对CYR30的抗性由1对显性基因控制,暂命名为Yrzhong21。该基因与位于小麦5AL染色体上的10个SSR位点连锁。与已定位于5A染色体上的抗条锈病基因的比较表明,Yrzhong21可能是一个抗条锈病的新基因。用侧翼标记(?)Ygwm186和IXbarc165检测中梁系列品种,其中仅17%扩增到与中梁21相同的位点,表明该基因在抗条锈病育种中可能有很大的应用潜力。
7.采用生理小种CYR31和Su11-11对三属麦1号进行遗传分析,结果表明三属麦1号对CYR31的抗病性是由一对显性基因控制,暂命名为YrS1；对Su11-11的抗病性是由一对隐性基因控制,暂命名为YrS2。利用BSA法对SSR引物进行筛选,获得了与YrS1连锁的7对SSR标记,并将YrS1定位于小麦3DS染色体；4对SSR标记与YrS2连锁,并将YrS2定位于小麦4DL染色体。绘制等的YrS1和YrS2遗传连锁图,将有助于分子选择辅助育种以及将该基因与其它抗病基因结合培育持久抗病品种。
8.通过对农家品种白大头进行遗传分析,结果表明白大头含有成株抗条锈基因。4个SSR标记和2个SRAP标记构建了与成株抗病基因YrBai连锁的遗传连锁图,该基因位于染色体6DL染色体上,这将有助于对白大头中该基因的合理和有效地利用。
9.通过对兰天1号苗期温室高温条件下的抗病性鉴定,以及F2群体遗传分析表明兰天1号对CYR32是由1对显性基因控制的,暂命名为YrLT1。利用BSA法对SSR引物和RGAP引物进行筛选,获得了与高温抗病基因YrLT1连锁的5对SSR标记和3对RGAP标记,并将YrLT1定位于小麦2DL染色体。这些标记将有助于将YrLT1导入其他小麦品种或者与其它抗病基因聚合培育持久抗病品种。
Stripe rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks, is one of the most important diseases of wheat worldwide. The best strategy of controlling stripe rust is to grow resistant cultivars. Two major types of resistance can be classified into seedling (also known as all-stage) resistance and adult plant resistance. Seedling resistance is usually race specific, but often provides complete control when it confers resistance to the prevalent races. Adult plant resistance is usually non-race specific and often requires high temperatures to function. One type of adult plant resistance is high-temperature adult-plant (HTAP) resistance. HTAP resistance, which is expressed in adult stages of plant development at higher temperatures, is non-race specific and durable Identification and excavation new stripe rust resistance genes is an important method for controlling stripe rust. Geneticists and plant breeders pay more attention to wild relatives and elite cultivars carrying durable resistance in order to search new resistant germplasms.
In this study, the wheat-P. huashanica Keng translocation lines H9014-14-4-6-1, H9015-17-1-9-6, H9014-121-5-5-9and varieties Zhongliang12, Zhongliang16, Zhongliang21, Sanshumai1, Baidatou and Lantian1were genetic analysis of stripe rust resistance genes and molecular mapping related resistance genes. The results are as follows:
1. The seedlings of the parents and F1plants, F2, F3and BC1generations were tested with Pst races under controlled greenhouse conditions. Two genes for resistance to stripe rust were identified, one dominant gene conferred resistance to Sull-4, temporarily designated YrH9014and the other recessive gene conferred resistance to CYR33. The bulked segregant analysis and simple sequence repeat (SSR) markers were used to identify polymorphic markers associated with YrH9014. Seven polymorphic SSR markers were used to genotype the F2population inoculated with Sull-4. A linkage map was constructed according to the genotypes of seven SSR markers and resistance gene. Based on the position of SSR marker, the resistance gene YrH9014was located on chromosome arm2BS. Amplification of a set of nulli-tetrasomic Chinese Spring lines with SSR marker Xbarc13indicated that YrH9014was located on chromosome2B. Based on chromosomal location, the reaction patterns and pedigree analysis, YrH9014should be a novel resistance gene to stripe rust.
2. To identify genes for the stripe rust resistances, H9015-17-1-9-6was crossed with Mingxian169. Genetic analysis of H9015-17-1-9-6indicated that the resistance to race CYR32was controlled by a single dominant gene, temporarily designated as YrH9015. Markers combined with BSA revealed that seven RGAP markers and three SSR markers located on chromosome5DL were linked to YrH9015. Pedigree analysis of H9015-17-1-9-6combined with SSR markers indicated that the highly resistance gene YrH9015derived from P. huashanica Keng, is a novel gene for resistance to stripe rust in wheat.
3. To determine the inheritence and map the resistance gene, segregating populations were developed from the cross between H9014-121-5-5-9and the susceptible cultivar Mingxian169. The seedlings of the F1, F2, and F2:3generations were tested with race CYR31. The results showed that the resistance in H9014-121-5-5-9was conferred by a single dominant gene. BSA and SSR markers were used to identify polymorphic markers associated with the resistance gene locus. Seven polymorphic SSR markers were linked to the resistance gene. A linkage map was constructed according to the genotypes of seven SSR markers and resistance gene. Based on the SSR marker position on wheat chromosome, the resistance gene was assigned on chromosome1AL, temporarily designated YrH9014. Based on chromosomal location, the reaction patterns and pedigree analysis, YrH9014should be a novel resistance gene to stripe rust. The molecular markers of the new resistance gene in H9014-121-5-5-9could be useful for marker-assisted selection (MAS) in breeding programs for against stripe rust.
4. Genetic analysis of Zhongliang12showed that resistance genes conferring to races CYR30,CYR31were all controlled by a single dominant gene, temporarily designated as YrZhongl2-1and YrZhongl2-2. Four SSR markers on chromosome7AL were linked to YrZhongl2-1.The SSR markers were Xwmc695、Xcfd20、Xbarc121、Xbarc49. And four SSR markers on chromosome1AL were linked to YrZhongl2-2. The SSR markers were Xpsp3003、Xcfd2129、Xwmc673、Xwmc51.
5. Inheritance of Zhongliang16by artificial inoculation with races CYR29and CYR30at seedlings showed that resistance genes conferring to races CYR29, CYR30were all controlled by a single dominant gene, temporarily designated as YrZhongl6-1and YrZhongl6-2. Five SSR markers on chromosome2AS were linked to YrZhongl6-1. And Four SSR markers on chromosome7BL were linked to YrZhongl6-2.
6. Seedlings of the F1, F2, and BC1generations from the cross between Zhongliang21and Mingxian169, as well as the parents, were tested with Pst race CYR30. The results showed that the stripe rust resistance in Zhongliang21was conferred by a single dominant gene, which was designated Yrzhong21, temporarily. Ten SSR markers located on chromosome arm5AL were linked to Yrzhong21with the nearest flanking markers of Xgwml86and Xbarc165. Based on chromosomal location, reactions to various pathotypes and pedigree analysis, we deduced that Yrzhong21was a novel resistance gene to stripe rust. Eighteen cultivars (lines) of Zhongliang series were tested with the flanking markers Xgwm186and Xbarcl65, and only17%showed the same banding pattern as that in Zhongliang21. This result suggested that17%of Zhongliang cultivars (lines) might carry Yrzhong21, indicating that this resistance gene has a potential application in wheat breeding program for strip rust resistance.
7. Genetic analysis of Sanshumai1demonstrated that resistance gene conferring to race CYR31were all controlled by a recessive gene, temporarily designated as YrS1; Resistance of Sanshumai1conferring to Sull-11was controlled by one dominant gene, temporarily designated as YrS2. Seven SSR markers on chromosome3DS were linked to YrS1. And Four SSR markers on chromosome4DL were linked to YrS2. These flanking markers got from this study should be useful for MAS in breeding programs for stripe rust.
8. To identify genes for the stripe rust resistances, Baidatou was crossed with Mingxian169. The result of genetic analysis showed Chinese wheat cultivar Baidatou has adult plant resistant to stripe rust. Two SRAP markers and four SSR markers were constructed adult pant resistance gene YrBai linkage map, which is located on the long arm of chromosome6D. Molecular markers should be useful in marker-assisted selection to incorporate these genes into commercial cultivars and combining them with other resistance genes for durable resistance.
9. Seedlings of the parents, and F1, F2and F2:3progeny were tested with races CYR32of Pst under controlled greenhouse conditions. Lantian1has a single partially dominant gene conferred resistance to race CYR32, designated as YrLT1. A linkage group of three RGAP markers and five SSR markers was constructed for YrLT1using166F2plants. Based on the marker consensus map and the position on wheat chromosome, the resistance gene was assigned on chromosome2DL. Amplification of a set of nulli-tetrasomic Chinese Spring lines with RGAP marker M3confirmed that the resistance gene was located on the long arm of chromosome2D. Because of its chromosomal location and the high-temperature resistance, this gene is different from previously described genes. This new gene and flanking markers should be useful in developing wheat cultivars with high level and possible durable resistance to stripe rust.
本研究对普通小麦-华山新麦草易位系H9014-14-4-6-1、H9015-17-1-9-6、H9014-121-5-5-9、中梁12、中梁16、中梁21、三属麦1号和兰天1号抗条锈病基因进行了遗传分析以及分子作图,取得了以下结果：
1.对普通小麦-华山新麦草易位系H9014-14-4-6-1与铭贤169杂交的F1、F2和F3代在苗期温室条件下进行遗传分析,结果表明H9014-14-4-6-1对Sul1-4的抗病性由一对显性基因控制,暂命名为YrH9014；对CYR33的抗病性由一对隐形基因控制。利用群体分离分析法(BSA)筛选到与抗病基因YrH9014连锁的简单重复序列(SSR)标记。用中国春缺四体进一步确定了SSR标记所在染色体的具体位置,抗锈基因YrH9014被定位在2B短臂上。对F2分离群体的142个单株进行分析,构建了与YrH9014连锁的遗传图谱。系谱分析结合分子标记结果表明,YrH9014可能是来自于华山新麦草,并与已知抗条锈病基因不同的。
2.通过对H9015-17-1-9-6与铭贤169杂交的F1、F2和F3代进行遗传分析,表明H9015-17-1-9-6对CYR32的抗病性由一对显性基因控制,暂命名为YrH9015。利用BSA法对RGAP引物和SSR引物进行筛选,获得了与抗条锈病基因YrH9015连锁的7对RGAP标记和3对SSR标记,并将YrH9015定位于小麦5DL染色体。YrH9015可能来自于华山新麦草,并与已知抗条锈病基因不同。
3.对H9014-121-5-5-9与铭贤169杂交的F1、F2和F3代进行遗传分析,结果表明H9014-121-5-5-9对CYR31的抗病性由一对显性基因控制,暂命名为YrHA。利用BSA法对SSR引物进行筛选,获得了与YrHA连锁的7对SSR标记,并将YrHA定位于小麦1AL染色体。系谱分析结合分子标记结果表明,YrHA可能来自于华山新麦草,并与已知抗条锈病基因不同。
4.经典遗传学分析表明,中梁12对CYR30和CYR31抗病性分别由一对显性基因控制,分别暂命名为YrZhong12-1和YrZhong12-2。BSA法结合SSR标记分析,7AL染色体上的Xwmc695、Xcfd20、Xbarc121、Xbarc49与YrZhong12-1连锁；1AL染色体上的Xpsp3003、Xcfd2129、Xwmc673、Xwmc51与YrZhong12-2连锁。
5.采用生理小种CYR29和CYR30对中梁16进行遗传分析,结果表明中梁16对CYR29和CYR30抗病性分别由一对显性基因控制,分别暂命名为YrZhong16-1和YrZhong16-2。利用BSA法对SSR引物进行筛选,获得了5个SSR标记,并将YrZhong16-1定位在2AS染色体上；5个SSR标记与YrZhong16-2连锁,并将YrZhong16-2定位在7BL染色体上。
6.采用CYR30小种对中梁21进行遗传分析,并利用SSR分子标记进行遗传作图,发现中梁21对CYR30的抗性由1对显性基因控制,暂命名为Yrzhong21。该基因与位于小麦5AL染色体上的10个SSR位点连锁。与已定位于5A染色体上的抗条锈病基因的比较表明,Yrzhong21可能是一个抗条锈病的新基因。用侧翼标记(?)Ygwm186和IXbarc165检测中梁系列品种,其中仅17%扩增到与中梁21相同的位点,表明该基因在抗条锈病育种中可能有很大的应用潜力。
7.采用生理小种CYR31和Su11-11对三属麦1号进行遗传分析,结果表明三属麦1号对CYR31的抗病性是由一对显性基因控制,暂命名为YrS1；对Su11-11的抗病性是由一对隐性基因控制,暂命名为YrS2。利用BSA法对SSR引物进行筛选,获得了与YrS1连锁的7对SSR标记,并将YrS1定位于小麦3DS染色体；4对SSR标记与YrS2连锁,并将YrS2定位于小麦4DL染色体。绘制等的YrS1和YrS2遗传连锁图,将有助于分子选择辅助育种以及将该基因与其它抗病基因结合培育持久抗病品种。
8.通过对农家品种白大头进行遗传分析,结果表明白大头含有成株抗条锈基因。4个SSR标记和2个SRAP标记构建了与成株抗病基因YrBai连锁的遗传连锁图,该基因位于染色体6DL染色体上,这将有助于对白大头中该基因的合理和有效地利用。
9.通过对兰天1号苗期温室高温条件下的抗病性鉴定,以及F2群体遗传分析表明兰天1号对CYR32是由1对显性基因控制的,暂命名为YrLT1。利用BSA法对SSR引物和RGAP引物进行筛选,获得了与高温抗病基因YrLT1连锁的5对SSR标记和3对RGAP标记,并将YrLT1定位于小麦2DL染色体。这些标记将有助于将YrLT1导入其他小麦品种或者与其它抗病基因聚合培育持久抗病品种。
Stripe rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks, is one of the most important diseases of wheat worldwide. The best strategy of controlling stripe rust is to grow resistant cultivars. Two major types of resistance can be classified into seedling (also known as all-stage) resistance and adult plant resistance. Seedling resistance is usually race specific, but often provides complete control when it confers resistance to the prevalent races. Adult plant resistance is usually non-race specific and often requires high temperatures to function. One type of adult plant resistance is high-temperature adult-plant (HTAP) resistance. HTAP resistance, which is expressed in adult stages of plant development at higher temperatures, is non-race specific and durable Identification and excavation new stripe rust resistance genes is an important method for controlling stripe rust. Geneticists and plant breeders pay more attention to wild relatives and elite cultivars carrying durable resistance in order to search new resistant germplasms.
In this study, the wheat-P. huashanica Keng translocation lines H9014-14-4-6-1, H9015-17-1-9-6, H9014-121-5-5-9and varieties Zhongliang12, Zhongliang16, Zhongliang21, Sanshumai1, Baidatou and Lantian1were genetic analysis of stripe rust resistance genes and molecular mapping related resistance genes. The results are as follows:
1. The seedlings of the parents and F1plants, F2, F3and BC1generations were tested with Pst races under controlled greenhouse conditions. Two genes for resistance to stripe rust were identified, one dominant gene conferred resistance to Sull-4, temporarily designated YrH9014and the other recessive gene conferred resistance to CYR33. The bulked segregant analysis and simple sequence repeat (SSR) markers were used to identify polymorphic markers associated with YrH9014. Seven polymorphic SSR markers were used to genotype the F2population inoculated with Sull-4. A linkage map was constructed according to the genotypes of seven SSR markers and resistance gene. Based on the position of SSR marker, the resistance gene YrH9014was located on chromosome arm2BS. Amplification of a set of nulli-tetrasomic Chinese Spring lines with SSR marker Xbarc13indicated that YrH9014was located on chromosome2B. Based on chromosomal location, the reaction patterns and pedigree analysis, YrH9014should be a novel resistance gene to stripe rust.
2. To identify genes for the stripe rust resistances, H9015-17-1-9-6was crossed with Mingxian169. Genetic analysis of H9015-17-1-9-6indicated that the resistance to race CYR32was controlled by a single dominant gene, temporarily designated as YrH9015. Markers combined with BSA revealed that seven RGAP markers and three SSR markers located on chromosome5DL were linked to YrH9015. Pedigree analysis of H9015-17-1-9-6combined with SSR markers indicated that the highly resistance gene YrH9015derived from P. huashanica Keng, is a novel gene for resistance to stripe rust in wheat.
3. To determine the inheritence and map the resistance gene, segregating populations were developed from the cross between H9014-121-5-5-9and the susceptible cultivar Mingxian169. The seedlings of the F1, F2, and F2:3generations were tested with race CYR31. The results showed that the resistance in H9014-121-5-5-9was conferred by a single dominant gene. BSA and SSR markers were used to identify polymorphic markers associated with the resistance gene locus. Seven polymorphic SSR markers were linked to the resistance gene. A linkage map was constructed according to the genotypes of seven SSR markers and resistance gene. Based on the SSR marker position on wheat chromosome, the resistance gene was assigned on chromosome1AL, temporarily designated YrH9014. Based on chromosomal location, the reaction patterns and pedigree analysis, YrH9014should be a novel resistance gene to stripe rust. The molecular markers of the new resistance gene in H9014-121-5-5-9could be useful for marker-assisted selection (MAS) in breeding programs for against stripe rust.
4. Genetic analysis of Zhongliang12showed that resistance genes conferring to races CYR30,CYR31were all controlled by a single dominant gene, temporarily designated as YrZhongl2-1and YrZhongl2-2. Four SSR markers on chromosome7AL were linked to YrZhongl2-1.The SSR markers were Xwmc695、Xcfd20、Xbarc121、Xbarc49. And four SSR markers on chromosome1AL were linked to YrZhongl2-2. The SSR markers were Xpsp3003、Xcfd2129、Xwmc673、Xwmc51.
5. Inheritance of Zhongliang16by artificial inoculation with races CYR29and CYR30at seedlings showed that resistance genes conferring to races CYR29, CYR30were all controlled by a single dominant gene, temporarily designated as YrZhongl6-1and YrZhongl6-2. Five SSR markers on chromosome2AS were linked to YrZhongl6-1. And Four SSR markers on chromosome7BL were linked to YrZhongl6-2.
6. Seedlings of the F1, F2, and BC1generations from the cross between Zhongliang21and Mingxian169, as well as the parents, were tested with Pst race CYR30. The results showed that the stripe rust resistance in Zhongliang21was conferred by a single dominant gene, which was designated Yrzhong21, temporarily. Ten SSR markers located on chromosome arm5AL were linked to Yrzhong21with the nearest flanking markers of Xgwml86and Xbarc165. Based on chromosomal location, reactions to various pathotypes and pedigree analysis, we deduced that Yrzhong21was a novel resistance gene to stripe rust. Eighteen cultivars (lines) of Zhongliang series were tested with the flanking markers Xgwm186and Xbarcl65, and only17%showed the same banding pattern as that in Zhongliang21. This result suggested that17%of Zhongliang cultivars (lines) might carry Yrzhong21, indicating that this resistance gene has a potential application in wheat breeding program for strip rust resistance.
7. Genetic analysis of Sanshumai1demonstrated that resistance gene conferring to race CYR31were all controlled by a recessive gene, temporarily designated as YrS1; Resistance of Sanshumai1conferring to Sull-11was controlled by one dominant gene, temporarily designated as YrS2. Seven SSR markers on chromosome3DS were linked to YrS1. And Four SSR markers on chromosome4DL were linked to YrS2. These flanking markers got from this study should be useful for MAS in breeding programs for stripe rust.
8. To identify genes for the stripe rust resistances, Baidatou was crossed with Mingxian169. The result of genetic analysis showed Chinese wheat cultivar Baidatou has adult plant resistant to stripe rust. Two SRAP markers and four SSR markers were constructed adult pant resistance gene YrBai linkage map, which is located on the long arm of chromosome6D. Molecular markers should be useful in marker-assisted selection to incorporate these genes into commercial cultivars and combining them with other resistance genes for durable resistance.
9. Seedlings of the parents, and F1, F2and F2:3progeny were tested with races CYR32of Pst under controlled greenhouse conditions. Lantian1has a single partially dominant gene conferred resistance to race CYR32, designated as YrLT1. A linkage group of three RGAP markers and five SSR markers was constructed for YrLT1using166F2plants. Based on the marker consensus map and the position on wheat chromosome, the resistance gene was assigned on chromosome2DL. Amplification of a set of nulli-tetrasomic Chinese Spring lines with RGAP marker M3confirmed that the resistance gene was located on the long arm of chromosome2D. Because of its chromosomal location and the high-temperature resistance, this gene is different from previously described genes. This new gene and flanking markers should be useful in developing wheat cultivars with high level and possible durable resistance to stripe rust.
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