苹果黑星病菌及斑点落叶病菌的寄主应答反应研究
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摘要
苹果黑星病和苹果斑点落叶病是苹果产业中较为严重的两种真菌病害，其致病菌因为营养型的不同对苹果的危害方式不同，而目前关于苹果对这两种病害的抗病系统却并不清楚。本论文主要目的是通过对于苹果与两类病原菌互作过程中苹果在不同水平的响应进行研究，获得了苹果对这两种营养型不同的病害的抗性反应相关信息，为苹果今后的抗病育种工作提供新的思路和理论基础。
一、苹果黑星病的研究
获得了苹果黑星病的抗性基因HcrVf2的转基因旭(McIntosh)株系，并分别对该转基因抗病株系、感病品种旭及抗病品种自由(Liberty)在转录水平和蛋白质组学水平上进行定量及半定量研究，获得的结果：
1、提高了苹果转基因效率，达到了20%的转化率，为功能基因验证及同源转基因的实现奠定了基础。
2、通过对HcrVf2转基因株系的温室接种试验研究，发现了对传统的R基因典型抗性反应不同的抗性表型并提出了合理的假设，其转录水平和蛋白质组水平上的初步研究结果，为“基因对基因”抗性模型的进一步完善提供了新的思路和实践基础。
二、苹果斑点落叶病的研究
建立苹果斑点落叶病离体接种体系，通过对110株杂交后代进行SSR分析和接种试验，挑选出抗病品种和感病品种，通过对病菌侵染后的显微观察、生长调节物质、蛋白质组学等水平的一系列研究，获得的结果：
1、显微观察发现寄主叶细胞栅栏组织结构与其抗病性密切相关，苹果叶片解剖结构特征可以作为鉴定苹果株系抗病性的一种快速手段，或苹果抗病性早期选择的辅助手段。
2、通过对内源激素的系统变化动态研究，发现ZR、IAA、ABA与多胺类内源激素与苹果抗病机制存在相关性，提出死体寄生营养型的苹果斑点落叶病的抗性机制不符合传统的基因对基因模型的假设。
3、蛋白组学研究共获得25个差异表达蛋白质点，有20个蛋白质点得到成功鉴定，其中11个蛋白分别参与苹果叶片细胞的光合作用、能量代谢、胁迫及防御反应。9个蛋白为蛋白质合成相关或功能未知蛋白。
综上所述，除了基因对基因模型中对植物的抗病反应起关键作用的抗病基因，对于基因背景复杂的苹果来说，相关的生理途径和代谢循环在抗病互作中同样十分重要。此外，根据苹果对黑星病和斑点落叶病侵染后的寄主响应研究，发现了抗苹果黑星病的非典型基因对基因反应和斑点落叶病的非基因对基因抗病模式的存在，并通过相关研究获得了新的思路和进展，为今后进一步开展苹果抗性育种工作奠定了理论和实践基础。
Alternaria alternata apple pathotype and Venturia inaequalis are two kinds of severest fungi diseasefor apple production, however the damages they cause are different as they have different habitat. Seriesof studies on the interaction between apple and two pathogens were processed to get more knowledgeand data for related applied and theoretical work in future.
Study on Veturia inaequalis
Intragenic lines with HcrVf2were transformed, and hemiquantitative and quantitative analysis wasapplied for transcript and proteomics in the process of infection for susceptible, resistant and transgenicresistant plants.
1. Transformation efficiency was improved by optimizing the protocol and media receipt whichprovide a chance for further gene functional evaluation and steping forward to cisgenic plant.
2. HcrVf2resistant symptom was observed being different from classic R gene symptom bygreenhouse inoculation test, which supplied new idea and foundament for improvement of “gene forgene” model.
Study on Alternaria alternata apple pathotype
A stable and efficient detached inoculation system was established. Susceptible and resistant varietieswere selected by inoculation and SSR test from100progeny group, furthermore microscopicobservation, proteomics and plant growth regulator determination were applied.
1. Leaf spongy cells supplied major nutrient for pathogenic infection, and the structure of palisadecells is closely related to disease resistance. Moreover the structure trait of palisade cells can be used asan assisted selection for resistance selection.
2、Systemic dynamics of endogenous hormones through interaction were studied, and thehypothesis on unfitness for “gene for gene” model of resistance mechanism to Alternaria alternataapple pathotype was innovatively proposed. Moreover the relationships between different endogenoushormones and resistance mechanism were found.
3、25differential expressed protein spots were gained by proteomics approach, in which20proteinspots were identified. Among those,11proteins were involved in photosynthesis, energy metabolismand defence response. Particularly,9proteins were still unknown, whose function was speculatedthough, playing a role of protein synthesis. Another3PR protein were inferred being defence-relatedprotein, which were deserved to be studied further.
To conclude, resistant gene plays a vital role for resistance response in “gene for gene” model.However related physiological, signal pathway and metabolic cycle are proved being also important forapple disease resistant interaction because of apple complicated genome background
一、苹果黑星病的研究
获得了苹果黑星病的抗性基因HcrVf2的转基因旭(McIntosh)株系，并分别对该转基因抗病株系、感病品种旭及抗病品种自由(Liberty)在转录水平和蛋白质组学水平上进行定量及半定量研究，获得的结果：
1、提高了苹果转基因效率，达到了20%的转化率，为功能基因验证及同源转基因的实现奠定了基础。
2、通过对HcrVf2转基因株系的温室接种试验研究，发现了对传统的R基因典型抗性反应不同的抗性表型并提出了合理的假设，其转录水平和蛋白质组水平上的初步研究结果，为“基因对基因”抗性模型的进一步完善提供了新的思路和实践基础。
二、苹果斑点落叶病的研究
建立苹果斑点落叶病离体接种体系，通过对110株杂交后代进行SSR分析和接种试验，挑选出抗病品种和感病品种，通过对病菌侵染后的显微观察、生长调节物质、蛋白质组学等水平的一系列研究，获得的结果：
1、显微观察发现寄主叶细胞栅栏组织结构与其抗病性密切相关，苹果叶片解剖结构特征可以作为鉴定苹果株系抗病性的一种快速手段，或苹果抗病性早期选择的辅助手段。
2、通过对内源激素的系统变化动态研究，发现ZR、IAA、ABA与多胺类内源激素与苹果抗病机制存在相关性，提出死体寄生营养型的苹果斑点落叶病的抗性机制不符合传统的基因对基因模型的假设。
3、蛋白组学研究共获得25个差异表达蛋白质点，有20个蛋白质点得到成功鉴定，其中11个蛋白分别参与苹果叶片细胞的光合作用、能量代谢、胁迫及防御反应。9个蛋白为蛋白质合成相关或功能未知蛋白。
综上所述，除了基因对基因模型中对植物的抗病反应起关键作用的抗病基因，对于基因背景复杂的苹果来说，相关的生理途径和代谢循环在抗病互作中同样十分重要。此外，根据苹果对黑星病和斑点落叶病侵染后的寄主响应研究，发现了抗苹果黑星病的非典型基因对基因反应和斑点落叶病的非基因对基因抗病模式的存在，并通过相关研究获得了新的思路和进展，为今后进一步开展苹果抗性育种工作奠定了理论和实践基础。
Alternaria alternata apple pathotype and Venturia inaequalis are two kinds of severest fungi diseasefor apple production, however the damages they cause are different as they have different habitat. Seriesof studies on the interaction between apple and two pathogens were processed to get more knowledgeand data for related applied and theoretical work in future.
Study on Veturia inaequalis
Intragenic lines with HcrVf2were transformed, and hemiquantitative and quantitative analysis wasapplied for transcript and proteomics in the process of infection for susceptible, resistant and transgenicresistant plants.
1. Transformation efficiency was improved by optimizing the protocol and media receipt whichprovide a chance for further gene functional evaluation and steping forward to cisgenic plant.
2. HcrVf2resistant symptom was observed being different from classic R gene symptom bygreenhouse inoculation test, which supplied new idea and foundament for improvement of “gene forgene” model.
Study on Alternaria alternata apple pathotype
A stable and efficient detached inoculation system was established. Susceptible and resistant varietieswere selected by inoculation and SSR test from100progeny group, furthermore microscopicobservation, proteomics and plant growth regulator determination were applied.
1. Leaf spongy cells supplied major nutrient for pathogenic infection, and the structure of palisadecells is closely related to disease resistance. Moreover the structure trait of palisade cells can be used asan assisted selection for resistance selection.
2、Systemic dynamics of endogenous hormones through interaction were studied, and thehypothesis on unfitness for “gene for gene” model of resistance mechanism to Alternaria alternataapple pathotype was innovatively proposed. Moreover the relationships between different endogenoushormones and resistance mechanism were found.
3、25differential expressed protein spots were gained by proteomics approach, in which20proteinspots were identified. Among those,11proteins were involved in photosynthesis, energy metabolismand defence response. Particularly,9proteins were still unknown, whose function was speculatedthough, playing a role of protein synthesis. Another3PR protein were inferred being defence-relatedprotein, which were deserved to be studied further.
To conclude, resistant gene plays a vital role for resistance response in “gene for gene” model.However related physiological, signal pathway and metabolic cycle are proved being also important forapple disease resistant interaction because of apple complicated genome background
引文
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