水稻广谱抗稻瘟病基因的遗传分析与定位
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摘要
稻瘟病是由子囊菌Magnaporthe grisea(Hebert)Barr[无性世代为Pyricularia grisea(Cooke)Sacc．]引起的，广泛分布于水稻栽培的国家和地区，是水稻上最重要的病害之一。寻找利用具有广谱抗性的抗源和抗性基因是当前控制稻瘟病的关键措施。谷梅4号是一个来自四川的地方品种，经过10多年的不同地区的种植试验，表现出广谱持久的抗瘟性，在实验室接种来自不同地区和国家的30个强致病力菌株，其抗29个，表现出广谱的抗瘟性，其抗谱明显强于现在认为具有广谱抗性的Pi-1、Pi-2、Pi-3等。
谷梅4号与两个感病粳稻Cpslo17和Maratelli杂交F_1，自交F_2，分别与感病亲本回交获得BC_1F_1，BC_1F_1自交获得BC_1F_2，然后通过接种稻瘟病菌株CH109、CH199、CH63、101／1／1、101／4／8、CH174、CH131进行遗传分析，发现谷梅4号对这些菌株的抗性均由一对显性单基因控制，对CH109接种的所有的367株F2群体移至田间，在分蘖盛期再注射接种菌株CH174、CH131，接种的抗感结果与CH109接种一致。这表明谷梅4号对这三个菌株的抗性为同一个基因，将此基因暂时定名为Pi-gm(t)。
对CH109接种的抗感分离群体，利用BSA分析法，以均匀分布在水稻12条染色体上105对SSR引物，把抗瘟基因Pi-gm(t)初步定位在第六条染色体的SRF5附近，然后在继续设计了一系列SSR标记和CAPS标记将Pi-gm(t)定位在引物C26348和C15959之间的5.1 cM范围内。利用距离Pi-gm(t)最近的标记C26348检测用不同菌株接种的定位群体构建的不同抗感池的多态性，它们的带型与CH109接种分离的抗感池的带型一致，这就表明Pi-gm(t)是一个广谱抗病基因或者其所处的区域存在紧密连锁的抗病基因簇。
在第6染色体的Pi-gm(t)基因附近还存在Pi-2、Pi-9、Pi-25，但接种定位群体的菌株CH109对Pi-2是有毒性的，所以可推测Pi-gm(t)和Pi-2可能不是同一个基因或存在某种差异，具体的关系还要经过Pi-gm(t)与Pi-2和Pi-9的等位性测定来进一步确定，与Pi-25定位接种菌株不同，其关系也不确定。
Rice blast, caused by the fungus Magnaporthe grisea, is one of the most serious and widespread diseases of rice (Oryza sativa L.) all over the world. In China, rice blast is considered as one of the most harmful diseases. To identify and use the broad-spectrum or durable resistance gene(s) is consided very important and effective way to control rice blast disease. Gu-mei 4 (GM4), a native variety from Sichuan, China, was identified in different disease nurseries for more than 20 years, and was characterized of broad-spectrum and durable resistance to blast. GM4 was inoculated with 30 blast isolates collected from different rice regions of China, Thailand and French. The results showed that GM4 could resist 29 of 30 isolates tested. Also the resistance spectrum of GM4 is broader than that of gene Pi-1, Pi-2, Pi-3.
The F1, F2, BC1, BC1F2 from the crosses between GM4 and susceptible verieties Cpslo17, Maratellu Suyunuo were obtained. Both parents and partial populations were sprayed by 7 isolates CH109, CH199, CH63, CH174, CH13K 101/1/1, 101/4/8, respectively. The inoculation results showed that blast resistance of GM4 was controlled by a single dominant gene tentively named Pi-gm(t). The F2 segregation of GM4/Cpslol7 populations inoculated with CHI09 isolate was inoculated by injecting CHI74 and CHI31 isolates at tillering stage. The result was the same to CHI09. This indicate that the Pi-gm(t) is a broad-spectrum resistance gene that controlled the resistance to 3 blast isolates.
We constructed the mapping population of Pi-gm(t) and conducted mapping study with 105 SSR makers using DNA extracted from different resistance/susceptible pools inoculated with the isolates CHI09. The gene Pi-gm(t) was preliminary mapped onto chromosome 6 near marker SRF5,and then we further designed some SSR and CAPS marker, mapping Pi-gm(t) in the region approximately 5.1cM between CAPS marker C26348 and C15959 , The CAPS marker C26348 amplified in different pools produced the same pattern to CH109,this result show that resistance to different isolates in GM4 is controlled by Pi-gm(t) gene, which is broad-spectrum gene or gene cluster.
The gene Pi-gm(l) is near to Pi-2 ^ Pi-9 and Pi-25 on chromosome 6. But the isolates CHI09 is virulence to Pi-2 presented in the rice line C101A51,also the resistance spectrum of Pi-gm(t)is broader than that of Pi-2, Pi-gm(t) should not be the same gene as Pi-2.The relation between Pi-gmft)
and Pi-2, Pi-9 will be identified by allelic test.Yet the relation between Pi-gm(t) and Pi-25 was not be
identified for different inoculating isolates.
谷梅4号与两个感病粳稻Cpslo17和Maratelli杂交F_1，自交F_2，分别与感病亲本回交获得BC_1F_1，BC_1F_1自交获得BC_1F_2，然后通过接种稻瘟病菌株CH109、CH199、CH63、101／1／1、101／4／8、CH174、CH131进行遗传分析，发现谷梅4号对这些菌株的抗性均由一对显性单基因控制，对CH109接种的所有的367株F2群体移至田间，在分蘖盛期再注射接种菌株CH174、CH131，接种的抗感结果与CH109接种一致。这表明谷梅4号对这三个菌株的抗性为同一个基因，将此基因暂时定名为Pi-gm(t)。
对CH109接种的抗感分离群体，利用BSA分析法，以均匀分布在水稻12条染色体上105对SSR引物，把抗瘟基因Pi-gm(t)初步定位在第六条染色体的SRF5附近，然后在继续设计了一系列SSR标记和CAPS标记将Pi-gm(t)定位在引物C26348和C15959之间的5.1 cM范围内。利用距离Pi-gm(t)最近的标记C26348检测用不同菌株接种的定位群体构建的不同抗感池的多态性，它们的带型与CH109接种分离的抗感池的带型一致，这就表明Pi-gm(t)是一个广谱抗病基因或者其所处的区域存在紧密连锁的抗病基因簇。
在第6染色体的Pi-gm(t)基因附近还存在Pi-2、Pi-9、Pi-25，但接种定位群体的菌株CH109对Pi-2是有毒性的，所以可推测Pi-gm(t)和Pi-2可能不是同一个基因或存在某种差异，具体的关系还要经过Pi-gm(t)与Pi-2和Pi-9的等位性测定来进一步确定，与Pi-25定位接种菌株不同，其关系也不确定。
Rice blast, caused by the fungus Magnaporthe grisea, is one of the most serious and widespread diseases of rice (Oryza sativa L.) all over the world. In China, rice blast is considered as one of the most harmful diseases. To identify and use the broad-spectrum or durable resistance gene(s) is consided very important and effective way to control rice blast disease. Gu-mei 4 (GM4), a native variety from Sichuan, China, was identified in different disease nurseries for more than 20 years, and was characterized of broad-spectrum and durable resistance to blast. GM4 was inoculated with 30 blast isolates collected from different rice regions of China, Thailand and French. The results showed that GM4 could resist 29 of 30 isolates tested. Also the resistance spectrum of GM4 is broader than that of gene Pi-1, Pi-2, Pi-3.
The F1, F2, BC1, BC1F2 from the crosses between GM4 and susceptible verieties Cpslo17, Maratellu Suyunuo were obtained. Both parents and partial populations were sprayed by 7 isolates CH109, CH199, CH63, CH174, CH13K 101/1/1, 101/4/8, respectively. The inoculation results showed that blast resistance of GM4 was controlled by a single dominant gene tentively named Pi-gm(t). The F2 segregation of GM4/Cpslol7 populations inoculated with CHI09 isolate was inoculated by injecting CHI74 and CHI31 isolates at tillering stage. The result was the same to CHI09. This indicate that the Pi-gm(t) is a broad-spectrum resistance gene that controlled the resistance to 3 blast isolates.
We constructed the mapping population of Pi-gm(t) and conducted mapping study with 105 SSR makers using DNA extracted from different resistance/susceptible pools inoculated with the isolates CHI09. The gene Pi-gm(t) was preliminary mapped onto chromosome 6 near marker SRF5,and then we further designed some SSR and CAPS marker, mapping Pi-gm(t) in the region approximately 5.1cM between CAPS marker C26348 and C15959 , The CAPS marker C26348 amplified in different pools produced the same pattern to CH109,this result show that resistance to different isolates in GM4 is controlled by Pi-gm(t) gene, which is broad-spectrum gene or gene cluster.
The gene Pi-gm(l) is near to Pi-2 ^ Pi-9 and Pi-25 on chromosome 6. But the isolates CHI09 is virulence to Pi-2 presented in the rice line C101A51,also the resistance spectrum of Pi-gm(t)is broader than that of Pi-2, Pi-gm(t) should not be the same gene as Pi-2.The relation between Pi-gmft)
and Pi-2, Pi-9 will be identified by allelic test.Yet the relation between Pi-gm(t) and Pi-25 was not be
identified for different inoculating isolates.
引文
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112. Zeigler RS. Recombination in Magnaporthe grisea. Annu Rev Phytopathol 1998; 36: 249-275.