双生病毒卫星DNA的复制及介导的病毒—介体互惠机制研究
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摘要
双生病毒是世界范围内广泛发生的一类单链环状DNA病毒,已在多个国家和地区的作物上造成毁灭性灾害。为了更好地了解双生病毒的致病机理及病毒的危害流行,本论文围绕双生病毒卫星DNA的复制专化性的分子机制和双生病毒与介体烟粉虱间接互惠的机制开展了以下两个方面的研究：
1中国番茄黄曲叶病毒与烟草曲茎病毒卫星复制专化性的分子机制研究
中国番茄黄曲叶病毒(TYLCCNV)和烟草曲茎病毒(TbCSV)的卫星DNAP分子可以被异源辅助病毒稳定复制,但是当同源及异源的DNAβ共同侵染时,异源的DNAβ会在后期丢失,而同源DNAβ则在整个侵染过程中一直存在,说明卫星分子对辅助病毒具有复制专化性。为了了解TYLCCNV和TbCSV卫星复制专化性的分子机制,首先构建了各种置换相应片段的突变体侵染性克隆,通过接种试验发现DNAP基因组上270bp左右的LCR区段决定了复制专化性,并且这种复制专化性是由辅助病毒上的复制蛋白(Rep, AC1)介导的。为了更进一步精细的寻找复制专化性的决定位点,通过序列比较分析发现LCR片段中存在一个具有重复序列的结构域(RBM),而TYLCCNV和TbCSV的卫星DNAP的这个结构域存在着序列上的不同。通过一系列的体外生化试验发现辅助病毒的Rep与其同源的RBM的结合能力强于其与异源的RBM的结合能力。通过构建该结构域的置换或缺失突变体侵染性克隆并且接种本氏烟,证实了该结构域正是这两种病毒卫星DNAP复制专化性的决定位点,且该结构域是卫星DNAβ复制所必须的。进一步利用烟草细胞系,证实了RBM结构域对于卫星DNAP的复制效率具有决定性的作用。在此基础上,利用中国番茄曲叶病毒(ToLCCNV)及其卫星DNAβ,与TYLCCNV或TbCSV的卫星DNAβ接种植物,证实了RBM结构域决定复制专化性的机制在双生病毒中是普遍存在的。
2中国番茄黄曲叶病毒与介体烟粉虱的互惠机制研究
前期的研究表明烟粉虱能与其传播的TYLCCNV通过寄主植物建立互惠关系。我们发现TYLCCNV伴随的卫星TYLCCNB是双生病毒与介体烟粉虱建立互惠互作关系所必须的。通过表达谱测序和qRT-PCR发现TYLCCNV和TYLCCNB共同侵染能抑制烟草的茉莉酸(Jasmonic acid, JA)防御途径；利用病毒诱导的基因沉默(VIGS)或转基因下调JA途径有利于烟粉虱的存活,通过外源涂抹茉莉酸甲酯(MeJA)或转基因上调JA途径则不利于烟粉虱的存活；进一步发现TYLCCNB所编码的βC1蛋白是影响植物防御途径变化从而使得介体昆虫和病毒建立互惠关系的关键因子,从而揭示了植物防御信号途径介导的介体昆虫和病毒建立互惠关系的重要机制；我们还发现了TYLCCNV和TYLCCNB共同侵染和烟粉虱取食分别下调和上调了普通烟中萜类化合物合成途径的一个关键基因5-epi-aristolochene synthase (NtEAS)的表达；通过VIGS或转基因沉默NtEAS有利于烟粉虱的存活,揭示了烟草的次生代谢途径是介导介体昆虫和病毒互惠关系的重要途径。
The Geminiviridae is the largest family of plant viruses with circular, single-stranded DNA genomes that cause devastating diseases of economically important crops world-wide. To get a better insight into the pathogenesis and epidemics of geminiviruses, the molecular mechanism of begomovirus satellite DNA replication specificity and the mechanism of begomovirus-vector whitefly mutualism were studied.
1. The molecular mechanism of begomovirus satellite DNA replication specificity
Tomato yellow leaf curl China virus (TYLCCNV) and Tobacco curly shoot virus (TbCSV), both obtained from Yunnan Province, were each found to be associated with a distant species of betasatellite molecule. Both tomato yellow leaf curl China betasatellite (TYLCCNB) and tobacco curly shoot betasatellite (TbCSB) can be trans-replicated stably by the noncognate begomovirus. When either TYLCCNV or TbCSV was associated with both betasatellite molecules simultaneously, the cognate betasatellite became dominant and the noncognate betasatellite became undetectable at late stages of infection. In the present work, we showed that the interaction between the LCR region of betasatellite and Rep protein of begomovirus determined the replication specificity. Furthermore, utilizing biochemistry assays in vitro, we found a novel Rep binding motif (RBM) in the LCR region and proved that the binding activity of Rep protein to its cognate RBM is stronger than to its noncognate RBM. Infectivity tests using infectious clones of RBM deletion mutant showed the RBM was involved in betasatellite infection and replication specificity. Finally we inoculated the Tomato leaf curl China virus (ToLCCNV) and its associated betasatellite together with TYLCCNB or TbCSB and found that the noncognate betasatellite containing the same RBM was detected even in late stage of infection while same the noncognate betasatellite containing the different RBM was undetectable in late stage of infection, indicating that the molecular mechanism of betasatellite replication specificity was ubiquitous.
2The mechanism of Begomovirus-vector whitefly mutualism
The whitefly can establish mutualism with the begomovirus TYLCCNV via crop plants. Here, we show that TYLCCNV and betasatellite co-infection suppresses jasmonic acid defenses in the plant. Using VIGS assays and transgenic plants to impair or enhance defenses mediated by jasmonic acid in the plant enhances or depresses the performance of the whitefly. We further generate the βC1transgenic tobacco plants, and demonstrate that the pathogenicity factor βC1encoded in the betasatellite is responsible for the initiation of suppression on plant defenses and contributes to the realization of the virus-vector mutualism. Our findings reveal that a begomovirus pathogenicity factor can suppress the plant defences and provide a mechanism whereby reduced defences result in increased whitefly vector multiplication. Our study also provides an interesting model to study the plant-pathogen-vector interactions through an integration of ecological, physiological and molecular approaches.
We also found that TYLCCNV and betasatellite co-infection or whitefly infested down or up regulated a key gene NtEAS involved in the terpenoid synthesis pathway, respectively. VIGS or transgenic silence NtEAS enhances the performance of the whitefly. Our study demonstrates that virus infection depletes the terpenoid-mediated plant defense against whiteflies, thereby favoring vector-virus mutualism. Our findings suggest that plant terpenoids play a key role in shaping vector-pathogen relationships. This is an evidence of secondary metabolic pathway mediated virus-vector mutualism on tobacco.
1中国番茄黄曲叶病毒与烟草曲茎病毒卫星复制专化性的分子机制研究
中国番茄黄曲叶病毒(TYLCCNV)和烟草曲茎病毒(TbCSV)的卫星DNAP分子可以被异源辅助病毒稳定复制,但是当同源及异源的DNAβ共同侵染时,异源的DNAβ会在后期丢失,而同源DNAβ则在整个侵染过程中一直存在,说明卫星分子对辅助病毒具有复制专化性。为了了解TYLCCNV和TbCSV卫星复制专化性的分子机制,首先构建了各种置换相应片段的突变体侵染性克隆,通过接种试验发现DNAP基因组上270bp左右的LCR区段决定了复制专化性,并且这种复制专化性是由辅助病毒上的复制蛋白(Rep, AC1)介导的。为了更进一步精细的寻找复制专化性的决定位点,通过序列比较分析发现LCR片段中存在一个具有重复序列的结构域(RBM),而TYLCCNV和TbCSV的卫星DNAP的这个结构域存在着序列上的不同。通过一系列的体外生化试验发现辅助病毒的Rep与其同源的RBM的结合能力强于其与异源的RBM的结合能力。通过构建该结构域的置换或缺失突变体侵染性克隆并且接种本氏烟,证实了该结构域正是这两种病毒卫星DNAP复制专化性的决定位点,且该结构域是卫星DNAβ复制所必须的。进一步利用烟草细胞系,证实了RBM结构域对于卫星DNAP的复制效率具有决定性的作用。在此基础上,利用中国番茄曲叶病毒(ToLCCNV)及其卫星DNAβ,与TYLCCNV或TbCSV的卫星DNAβ接种植物,证实了RBM结构域决定复制专化性的机制在双生病毒中是普遍存在的。
2中国番茄黄曲叶病毒与介体烟粉虱的互惠机制研究
前期的研究表明烟粉虱能与其传播的TYLCCNV通过寄主植物建立互惠关系。我们发现TYLCCNV伴随的卫星TYLCCNB是双生病毒与介体烟粉虱建立互惠互作关系所必须的。通过表达谱测序和qRT-PCR发现TYLCCNV和TYLCCNB共同侵染能抑制烟草的茉莉酸(Jasmonic acid, JA)防御途径；利用病毒诱导的基因沉默(VIGS)或转基因下调JA途径有利于烟粉虱的存活,通过外源涂抹茉莉酸甲酯(MeJA)或转基因上调JA途径则不利于烟粉虱的存活；进一步发现TYLCCNB所编码的βC1蛋白是影响植物防御途径变化从而使得介体昆虫和病毒建立互惠关系的关键因子,从而揭示了植物防御信号途径介导的介体昆虫和病毒建立互惠关系的重要机制；我们还发现了TYLCCNV和TYLCCNB共同侵染和烟粉虱取食分别下调和上调了普通烟中萜类化合物合成途径的一个关键基因5-epi-aristolochene synthase (NtEAS)的表达；通过VIGS或转基因沉默NtEAS有利于烟粉虱的存活,揭示了烟草的次生代谢途径是介导介体昆虫和病毒互惠关系的重要途径。
The Geminiviridae is the largest family of plant viruses with circular, single-stranded DNA genomes that cause devastating diseases of economically important crops world-wide. To get a better insight into the pathogenesis and epidemics of geminiviruses, the molecular mechanism of begomovirus satellite DNA replication specificity and the mechanism of begomovirus-vector whitefly mutualism were studied.
1. The molecular mechanism of begomovirus satellite DNA replication specificity
Tomato yellow leaf curl China virus (TYLCCNV) and Tobacco curly shoot virus (TbCSV), both obtained from Yunnan Province, were each found to be associated with a distant species of betasatellite molecule. Both tomato yellow leaf curl China betasatellite (TYLCCNB) and tobacco curly shoot betasatellite (TbCSB) can be trans-replicated stably by the noncognate begomovirus. When either TYLCCNV or TbCSV was associated with both betasatellite molecules simultaneously, the cognate betasatellite became dominant and the noncognate betasatellite became undetectable at late stages of infection. In the present work, we showed that the interaction between the LCR region of betasatellite and Rep protein of begomovirus determined the replication specificity. Furthermore, utilizing biochemistry assays in vitro, we found a novel Rep binding motif (RBM) in the LCR region and proved that the binding activity of Rep protein to its cognate RBM is stronger than to its noncognate RBM. Infectivity tests using infectious clones of RBM deletion mutant showed the RBM was involved in betasatellite infection and replication specificity. Finally we inoculated the Tomato leaf curl China virus (ToLCCNV) and its associated betasatellite together with TYLCCNB or TbCSB and found that the noncognate betasatellite containing the same RBM was detected even in late stage of infection while same the noncognate betasatellite containing the different RBM was undetectable in late stage of infection, indicating that the molecular mechanism of betasatellite replication specificity was ubiquitous.
2The mechanism of Begomovirus-vector whitefly mutualism
The whitefly can establish mutualism with the begomovirus TYLCCNV via crop plants. Here, we show that TYLCCNV and betasatellite co-infection suppresses jasmonic acid defenses in the plant. Using VIGS assays and transgenic plants to impair or enhance defenses mediated by jasmonic acid in the plant enhances or depresses the performance of the whitefly. We further generate the βC1transgenic tobacco plants, and demonstrate that the pathogenicity factor βC1encoded in the betasatellite is responsible for the initiation of suppression on plant defenses and contributes to the realization of the virus-vector mutualism. Our findings reveal that a begomovirus pathogenicity factor can suppress the plant defences and provide a mechanism whereby reduced defences result in increased whitefly vector multiplication. Our study also provides an interesting model to study the plant-pathogen-vector interactions through an integration of ecological, physiological and molecular approaches.
We also found that TYLCCNV and betasatellite co-infection or whitefly infested down or up regulated a key gene NtEAS involved in the terpenoid synthesis pathway, respectively. VIGS or transgenic silence NtEAS enhances the performance of the whitefly. Our study demonstrates that virus infection depletes the terpenoid-mediated plant defense against whiteflies, thereby favoring vector-virus mutualism. Our findings suggest that plant terpenoids play a key role in shaping vector-pathogen relationships. This is an evidence of secondary metabolic pathway mediated virus-vector mutualism on tobacco.
引文
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