大豆疫霉菌无毒基因的克隆及马铃薯抗病品种紫花白抗晚疫病机制的研究
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摘要
卵菌纲是近些年来由鞭毛菌亚门中新分出的一大类，能够侵染很多重要的粮食作物、林业、观赏植物进而对自然生态系统造成严重影响。大豆疫霉病和马铃薯晚疫病是由卵菌纲真菌Phytophthora sojae和Phytophthora infestans分别侵染大豆和马铃薯后发生的两种主要病害。目前，防治这两种病害最为有效、经济以及环保的方式依然是培育优良的抗病品种。因此，利用分子生物学技术克隆和鉴定致病疫霉的无毒基因、解析马铃薯抗晚疫病机制可以为大豆和马铃薯的抗病育种奠定基础。
本论文中采用了筛选候选Avh基因方法克隆了P.sojae无毒基因Avr1d。利用遗传作图分析确定了Avr1d与Avh6共分离。在含有抗性基因Rps1d的大豆叶片中进行功能验证，证实了Avh6就是Avr1d。基因比对结果得知Avr1d基因在对Rps1d有毒的生理小种的基因组中是缺失的。在对11个大豆疫霉生理小种进行基因测序分析得知，Avr1d有2个allele。在Rps1d大豆植株中瞬时表达全长野生型Avr1d蛋白和全长RFLR突变体蛋白（RFLR突变为AAAA）的结果表明，RFLR保守区域不能介导Avr1d分泌到寄主细胞的过程中。脂类结合实验结果表明effector区域的正电荷氨基酸决定Avr1d蛋白与寄主细胞膜上的PtdIns(4)P结合。
论文中利用分离群体分组分析法结合新一代测序技术找到了与无毒基因Avr1c紧密连锁的3个单核苷酸多态性位点。利用MapMaker软件和生物信息学比对软件，最后确定Avh275c在遗传群体中的基因型与Avr1c的表型共分离。基因枪瞬时表达实验也同时证明Avh275c就是Avr1c。对15个无毒生理小种测序分析得知，Avr1c有3个allele。Avr1c与Avr1a在大豆疫霉基因组上的位置非常相近。这两个基因在氨基酸序列N端有高度的一致性，差异在C端显著较大。
通过对马铃薯抗感品种紫花白和夏坡蒂在接种晚疫病菌前后活性氧产生的速度和活性氧量的积累以及抗病信号路径中重要蛋白在转录水平的变化来阐述马铃薯抗病品种紫花白抗晚疫病菌的机制。结果表明，活性氧爆发是抗病品种紫花白抵抗晚疫病菌侵入重要的抗病信号机制。活性氧在抗病品种中积累主要通过定位于细胞膜上的NADPH氧化酶实现。同时，通过对主要的抗病信号路径中（SA和JA路径）重要蛋白接种后转录水平的检测发现信号分子也介入到了紫花白对晚疫病的抵抗过程中。
Oomycetes which are separated from mastigomycotina recently, can infect manyplants. It causes serious problems on natural ecosystems by infect crops, forestry andornamental plant. Soybean blight and potato late blight are two main diseases in soybeansand potatoes caused by Oomycetes fungi Phytophthora sojae and Phytophthora infestansseparately. Currently, the most effective, economic and environmentally friendly way ofprevention and treatment of these two diseases rely on cultivating resistant varieties.Therefore, cloning and identification of avirulence gene in Phytophthora using molecularbiology techniques and unravel the mechanism of potato resistance to late blight diseasecould lay the foundation for soybean and potato disease resistance breeding.
In this thesis, two P.sojae avirulence genes Avr1d was cloned by using candidate Avhgenes screening method. Genetic mapping of the candidate Avh genes in F2mappingpopulation suggested that Avh6co-segregates with Avr1d. Transient expression assaysconfirm that Avr1d triggers cell death specifically in Rps1d soybean plants. Alignmentbetween Avr1d and the whole genome shotgun sequencing from5P.sojae strains wereperformed. The results showed that Avr1d gene is present in P. sojae strains that areavirulent to host which contain Rps1d resistance gene, whereas the gene is deleted from thegenome of virulent strains. Two allelee of Avh6were identified in the avirulent strains andthey encoded slightly different protein products. Transient expression of full-length wildtype Avr1d and full-length Avr1d with RFLR (Arg-Phe-Leu-Arg) mutant to AAAA (Ala-Ala-Ala-Ala) showed that RFLR region could not lead the Avr1d protein into the host cells.The liposome binding experiment data indicated that Avr1d could bind to PtdIns (4) P andthe binding is achieved through positive amino acids of effector domain of Avr1d, insteadof by the RFLR motif.
Three Avr1c linked single nucleotide polymorphisms (SNPs) were found by BulkSegregate Analysis combined with next-generation sequencing technology. Genetic andphysical mapping results suggested Avh275c co-segregate with Avr1c. Transientexpression experiments also prove that Avh275c is Avr1c. Haplotype sequencing analysisshowed that three alleles of Avr1c exist in the strain collection. Previous work proved thatAvh275c located very close to Avr1a on P.sojae genome. Comparison of the sequences from these two gene suggested Avr1c and Avr1a encoded slightly different protein. Themost variable region of the protein was found within N terminal effector domain.
The infection of P.infestans on potato will trigger the host a series of resistanceresponds. To know the resistance mechanism of potato resistant cultivar Zihuabai,accumulation of reactive oxygen species (ROS), activities of oxygen scavenging enzymes,and transcription level of some resistance genes were determined in both resistant andsusceptible potato cultivars inoculated with P. infestans. Our results showed that ROSaccumulation is an important signal pathway that involved in the procedure of Zihuabairesistant to P.infestans. The accumulation of ROS in the resistant cultivar depends on themembrane NADPH oxidase. The results of transcript expression of important protein alsoindicated that signaling molecules also involved in regulating the resistance of potatocultivar Zihuabai to P. infestans.
本论文中采用了筛选候选Avh基因方法克隆了P.sojae无毒基因Avr1d。利用遗传作图分析确定了Avr1d与Avh6共分离。在含有抗性基因Rps1d的大豆叶片中进行功能验证，证实了Avh6就是Avr1d。基因比对结果得知Avr1d基因在对Rps1d有毒的生理小种的基因组中是缺失的。在对11个大豆疫霉生理小种进行基因测序分析得知，Avr1d有2个allele。在Rps1d大豆植株中瞬时表达全长野生型Avr1d蛋白和全长RFLR突变体蛋白（RFLR突变为AAAA）的结果表明，RFLR保守区域不能介导Avr1d分泌到寄主细胞的过程中。脂类结合实验结果表明effector区域的正电荷氨基酸决定Avr1d蛋白与寄主细胞膜上的PtdIns(4)P结合。
论文中利用分离群体分组分析法结合新一代测序技术找到了与无毒基因Avr1c紧密连锁的3个单核苷酸多态性位点。利用MapMaker软件和生物信息学比对软件，最后确定Avh275c在遗传群体中的基因型与Avr1c的表型共分离。基因枪瞬时表达实验也同时证明Avh275c就是Avr1c。对15个无毒生理小种测序分析得知，Avr1c有3个allele。Avr1c与Avr1a在大豆疫霉基因组上的位置非常相近。这两个基因在氨基酸序列N端有高度的一致性，差异在C端显著较大。
通过对马铃薯抗感品种紫花白和夏坡蒂在接种晚疫病菌前后活性氧产生的速度和活性氧量的积累以及抗病信号路径中重要蛋白在转录水平的变化来阐述马铃薯抗病品种紫花白抗晚疫病菌的机制。结果表明，活性氧爆发是抗病品种紫花白抵抗晚疫病菌侵入重要的抗病信号机制。活性氧在抗病品种中积累主要通过定位于细胞膜上的NADPH氧化酶实现。同时，通过对主要的抗病信号路径中（SA和JA路径）重要蛋白接种后转录水平的检测发现信号分子也介入到了紫花白对晚疫病的抵抗过程中。
Oomycetes which are separated from mastigomycotina recently, can infect manyplants. It causes serious problems on natural ecosystems by infect crops, forestry andornamental plant. Soybean blight and potato late blight are two main diseases in soybeansand potatoes caused by Oomycetes fungi Phytophthora sojae and Phytophthora infestansseparately. Currently, the most effective, economic and environmentally friendly way ofprevention and treatment of these two diseases rely on cultivating resistant varieties.Therefore, cloning and identification of avirulence gene in Phytophthora using molecularbiology techniques and unravel the mechanism of potato resistance to late blight diseasecould lay the foundation for soybean and potato disease resistance breeding.
In this thesis, two P.sojae avirulence genes Avr1d was cloned by using candidate Avhgenes screening method. Genetic mapping of the candidate Avh genes in F2mappingpopulation suggested that Avh6co-segregates with Avr1d. Transient expression assaysconfirm that Avr1d triggers cell death specifically in Rps1d soybean plants. Alignmentbetween Avr1d and the whole genome shotgun sequencing from5P.sojae strains wereperformed. The results showed that Avr1d gene is present in P. sojae strains that areavirulent to host which contain Rps1d resistance gene, whereas the gene is deleted from thegenome of virulent strains. Two allelee of Avh6were identified in the avirulent strains andthey encoded slightly different protein products. Transient expression of full-length wildtype Avr1d and full-length Avr1d with RFLR (Arg-Phe-Leu-Arg) mutant to AAAA (Ala-Ala-Ala-Ala) showed that RFLR region could not lead the Avr1d protein into the host cells.The liposome binding experiment data indicated that Avr1d could bind to PtdIns (4) P andthe binding is achieved through positive amino acids of effector domain of Avr1d, insteadof by the RFLR motif.
Three Avr1c linked single nucleotide polymorphisms (SNPs) were found by BulkSegregate Analysis combined with next-generation sequencing technology. Genetic andphysical mapping results suggested Avh275c co-segregate with Avr1c. Transientexpression experiments also prove that Avh275c is Avr1c. Haplotype sequencing analysisshowed that three alleles of Avr1c exist in the strain collection. Previous work proved thatAvh275c located very close to Avr1a on P.sojae genome. Comparison of the sequences from these two gene suggested Avr1c and Avr1a encoded slightly different protein. Themost variable region of the protein was found within N terminal effector domain.
The infection of P.infestans on potato will trigger the host a series of resistanceresponds. To know the resistance mechanism of potato resistant cultivar Zihuabai,accumulation of reactive oxygen species (ROS), activities of oxygen scavenging enzymes,and transcription level of some resistance genes were determined in both resistant andsusceptible potato cultivars inoculated with P. infestans. Our results showed that ROSaccumulation is an important signal pathway that involved in the procedure of Zihuabairesistant to P.infestans. The accumulation of ROS in the resistant cultivar depends on themembrane NADPH oxidase. The results of transcript expression of important protein alsoindicated that signaling molecules also involved in regulating the resistance of potatocultivar Zihuabai to P. infestans.
引文
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