一粒小麦中抗白粉病新基因的发掘
摘要
由白粉菌侵染引起的白粉病是小麦的重要病害之一，抗病品种的培育和利用是控制该病害的关键措施。为了发现新的小麦抗白粉病基因，对抗白粉病的8份一粒小麦和2份二粒小麦种质进行了抗性遗传分析。结果表明一粒小麦223410、223418、223423和223426携一个显性抗病基因；223404和223414携一个隐性抗病基因；一粒小麦223405、223409和二粒小麦223309、223310携多个抗病基因。等位性测验表明223410、223418、223423和223426的抗病基因相互等位或者紧密相斥连锁，223409中有一个抗病基因与223410的抗病基因相互等位或者紧密连锁，223404的抗病基因与223410的抗病基因在遗传上相互独立，223309与223310可能携有相互等位或者紧密连锁的抗病基因。
标记分析表明223410的抗病基因Mlm2033和223426的抗病基因Mlm80均与染色体7A长臂的Xgwm344紧密连锁。为了比较它们与位于7AL的Pml的关系，将已报道的与Pml紧密连锁的五个RFLP标记转化成了STS标记，其中三个在作图亲本间检测到多态，并被定位于紧靠Mlm2033与Mlm80的位置。其中来源于RFLP标记PSR680的Xmag2185与Mlm2033和Mlm80两基因的遗传距离不到2 cM。此外通过比较基因组作图和EST-STS标记分析还发现了与这两个抗病基因连锁的多个标记。该研究为Pml位点的图位克隆打下了基础，同时也将促进这些抗病基因向普通小麦转移。
标记分析表明一粒小麦223404的抗病基因mlm404位于染色体5AL，SSR标记位点Xzmf39和Xzmg43分别位于基因的两侧，与其遗传距离分别为2.8 cM和2.0 cM。为了精细定位该基因，我们用5AL的RFLP探针和EST开发的STS标记丰富了mlm404附近的标记密度，发现了与其更为紧密连锁的STS标记MAG1493。mlm404与大麦抗白粉病基因Mlo直系同源基因TmMlo紧密连锁，遗传距离为1.0 cM。与感病材料223389相比，抗病材料223404的TmMlo编码的蛋白有一个氨基酸替换。此外，223404的TmMlo表达受到抑制。基因mlm404是一个未报道的新抗白粉病基因位点。
遗传分析表明一粒小麦223405的白粉病抗性由一个主效隐性抗病基因加微效基因共同控制。利用177个F_(2：3)家系进行简单区间作图在染色体5AL的Xzmf39／Xmag1491～Xmag1493区间发现了一个主效QTL Qpm. nau-5A。该QTL解释了57.8％的表型变异，其抗性等位位点来源于抗病亲本223405。来自于感亲本223389的等位位点呈显性，显性度为73.4％。连锁的分子标记表明Qpm. nau-5A可能是mlm404的一个等位基因。
Powdery mildew, caused by Blumeria graminis(DC.) E. O. Speer f. sp. tritici (Bgt), isone of the devastating diseases of wheat. Breeding and deploying cultivars carryingresistance genes against powdery mildew is the key measure to control this disease. Tosearch for new powdery mildew resistance genes, eight einkorn accessions and two emmeraccessions were studied for the inheritance of their resistance to powdery mildew. Theresults showed that einkorn accessions 223410, 223418, 223423 and 223426 each carried adominant resistance gene, 223404 and 223414 each had a recessive resistance gene, andeinkorn accessions 223405 and 223409 and emmer accessions 22309 and 223310 eachcarried more than one resistance genes. Allelism tests indicated that the resistance genes in223410, 223418, 223423 and 223426 were likely allelic or tightly linked to each other, aresistance gene in 223409 was also likely allelic or linked to the resistance gene in 223410,the resistance gene in 223404 was inherited independent of that in 223410, and 223309might carry resistance genes allelic or linked to that in 223310.
Marker mapping of the resistance gene, Mlm2033, in 223410 and that, Mlm80, in223426 indicated that they were both closely linked to Xgwm344 on the long arm ofchromosome 7A. To establish their genetic relationship with Pml on 7AL, five RFLPmarkers previously reported to co-segregate with Pmla were converted to STS markers,three of which detected polymorphism between the mapping parents and were mappedclose to Mlm2033 or Mlm80. Xmag2185, the locus determined by the STS marker derivedfrom PSR680, one of the RFLP markers, was placed less than 2 cM away from Mlm2033and Mlm80, suggesting that they are allelic or tightly linked to Pmla. In addition, through comparative and EST mapping, more markers linked to these two genes were identified.This research established the basis for map-based cloning of the Pml locus. The markersfor both genes will also facilitate their transfer to wheat.
Marker mapping of the resistance gene, mlm404, in 223404 showed that it was flankedby SSR markers loci Xcfd39 and Xgwm126 on chromosome 5AL with a genetic distance of2.8 and 2.0 cM, respectively. To fine mapping the resistance gene, STS markers weredesigned using some RFLP probes and ESTs mapped on the long arm of chromsome 5ALto densify markers on the chromosome region surrounding mlm404, and the STS markermore closely linked to mlm404, MAG1493, was identified. The resistance gene mlm404was tightly linked to TmMlo, the barley Mlo ortholog in T. monococcum, with a geneticdistance 1.0 cM. One amino acid was substituted in the protein encoded by TmMlo of theaccession 223404 compared with that of the susceptible accession 2233389. In addition, thetranscription of TmMlo of 223404 was suppressed. Currently, no known powdery mildewresistance gene was mappd to the locu of mlm404.
Genetic analysis showed that a major gene plus several minor genes controlled thepowdery mildew resistance in 223405. A major QTL named as Qpm. nau-5A was identifiedthrough simple interval mapping using 177 F_(2:3) families in the intervalXzmf39/Xmag1491～Xmag1493 on chromosome 5AL. This QTL explained 57.8%phenotypic variation, of which the resistance allele originated from the resistant parent223405 and the dominant allele originated from susceptible parent 223389 with a degree of73.4%. Qpm. nau-5A was likely allelic to mlm404 according to its linked molecularmarkers.
标记分析表明223410的抗病基因Mlm2033和223426的抗病基因Mlm80均与染色体7A长臂的Xgwm344紧密连锁。为了比较它们与位于7AL的Pml的关系，将已报道的与Pml紧密连锁的五个RFLP标记转化成了STS标记，其中三个在作图亲本间检测到多态，并被定位于紧靠Mlm2033与Mlm80的位置。其中来源于RFLP标记PSR680的Xmag2185与Mlm2033和Mlm80两基因的遗传距离不到2 cM。此外通过比较基因组作图和EST-STS标记分析还发现了与这两个抗病基因连锁的多个标记。该研究为Pml位点的图位克隆打下了基础，同时也将促进这些抗病基因向普通小麦转移。
标记分析表明一粒小麦223404的抗病基因mlm404位于染色体5AL，SSR标记位点Xzmf39和Xzmg43分别位于基因的两侧，与其遗传距离分别为2.8 cM和2.0 cM。为了精细定位该基因，我们用5AL的RFLP探针和EST开发的STS标记丰富了mlm404附近的标记密度，发现了与其更为紧密连锁的STS标记MAG1493。mlm404与大麦抗白粉病基因Mlo直系同源基因TmMlo紧密连锁，遗传距离为1.0 cM。与感病材料223389相比，抗病材料223404的TmMlo编码的蛋白有一个氨基酸替换。此外，223404的TmMlo表达受到抑制。基因mlm404是一个未报道的新抗白粉病基因位点。
遗传分析表明一粒小麦223405的白粉病抗性由一个主效隐性抗病基因加微效基因共同控制。利用177个F_(2：3)家系进行简单区间作图在染色体5AL的Xzmf39／Xmag1491～Xmag1493区间发现了一个主效QTL Qpm. nau-5A。该QTL解释了57.8％的表型变异，其抗性等位位点来源于抗病亲本223405。来自于感亲本223389的等位位点呈显性，显性度为73.4％。连锁的分子标记表明Qpm. nau-5A可能是mlm404的一个等位基因。
Powdery mildew, caused by Blumeria graminis(DC.) E. O. Speer f. sp. tritici (Bgt), isone of the devastating diseases of wheat. Breeding and deploying cultivars carryingresistance genes against powdery mildew is the key measure to control this disease. Tosearch for new powdery mildew resistance genes, eight einkorn accessions and two emmeraccessions were studied for the inheritance of their resistance to powdery mildew. Theresults showed that einkorn accessions 223410, 223418, 223423 and 223426 each carried adominant resistance gene, 223404 and 223414 each had a recessive resistance gene, andeinkorn accessions 223405 and 223409 and emmer accessions 22309 and 223310 eachcarried more than one resistance genes. Allelism tests indicated that the resistance genes in223410, 223418, 223423 and 223426 were likely allelic or tightly linked to each other, aresistance gene in 223409 was also likely allelic or linked to the resistance gene in 223410,the resistance gene in 223404 was inherited independent of that in 223410, and 223309might carry resistance genes allelic or linked to that in 223310.
Marker mapping of the resistance gene, Mlm2033, in 223410 and that, Mlm80, in223426 indicated that they were both closely linked to Xgwm344 on the long arm ofchromosome 7A. To establish their genetic relationship with Pml on 7AL, five RFLPmarkers previously reported to co-segregate with Pmla were converted to STS markers,three of which detected polymorphism between the mapping parents and were mappedclose to Mlm2033 or Mlm80. Xmag2185, the locus determined by the STS marker derivedfrom PSR680, one of the RFLP markers, was placed less than 2 cM away from Mlm2033and Mlm80, suggesting that they are allelic or tightly linked to Pmla. In addition, through comparative and EST mapping, more markers linked to these two genes were identified.This research established the basis for map-based cloning of the Pml locus. The markersfor both genes will also facilitate their transfer to wheat.
Marker mapping of the resistance gene, mlm404, in 223404 showed that it was flankedby SSR markers loci Xcfd39 and Xgwm126 on chromosome 5AL with a genetic distance of2.8 and 2.0 cM, respectively. To fine mapping the resistance gene, STS markers weredesigned using some RFLP probes and ESTs mapped on the long arm of chromsome 5ALto densify markers on the chromosome region surrounding mlm404, and the STS markermore closely linked to mlm404, MAG1493, was identified. The resistance gene mlm404was tightly linked to TmMlo, the barley Mlo ortholog in T. monococcum, with a geneticdistance 1.0 cM. One amino acid was substituted in the protein encoded by TmMlo of theaccession 223404 compared with that of the susceptible accession 2233389. In addition, thetranscription of TmMlo of 223404 was suppressed. Currently, no known powdery mildewresistance gene was mappd to the locu of mlm404.
Genetic analysis showed that a major gene plus several minor genes controlled thepowdery mildew resistance in 223405. A major QTL named as Qpm. nau-5A was identifiedthrough simple interval mapping using 177 F_(2:3) families in the intervalXzmf39/Xmag1491～Xmag1493 on chromosome 5AL. This QTL explained 57.8%phenotypic variation, of which the resistance allele originated from the resistant parent223405 and the dominant allele originated from susceptible parent 223389 with a degree of73.4%. Qpm. nau-5A was likely allelic to mlm404 according to its linked molecularmarkers.
引文
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