水稻条纹病毒基因NS2致病作用机理分析
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摘要
水稻条纹叶枯病是水稻的重要病毒病之一，严重危害着我国的水稻生产，其病原为水稻条纹病毒(Rice stripe virus, RSV)，是纤细病毒属(Tenuivirus)的代表成员。近年随着生物技术的发展，对该病毒的研究也日趋深入，特别是两大模式植物即本氏烟和拟南芥被确认为可被RSV侵染后，使得对该病毒的研究可以克服在寄主水稻研究上的诸如水稻培养周期长、转基因植株培育时间久等种种障碍和困难。RSV编码的NS2蛋白是新发现的一个弱基因沉默抑制子，定位在细胞核，能与水稻基因沉默抑制子3(suppressor of gene silencing3, SGS3)互作，SGS3是植物内源基因，相对保守，研究表明拟南芥SGS3突变体表现出强感病性，但目前对NS2基因如何致病以及OsSGS3基因是否具有抗病性还未被深入研究。因此，本文利用病毒诱导基因沉默技术(Virus inducing gene silencing, VIGS)，在本氏烟上对NS2基因的致病性和OsSGS3基因的抗病性进行分析；同时构建了OsSGS3-RNAi转基因水稻，为研究其在RSV的抗病调控功能打下基础。
本文研究内容主要分为三部分。第一部分，通过共定位方法进一步确定NS2在细胞核内的具体位置，并通过双荧光互补实验(Bimolecular fluorescencecomplementation, BiFC)确定NS2与三种寄主即拟南芥、本氏烟和水稻的核仁和柯浩体的互作情况；第二部分，利用VIGS技术分别沉默基因NbFib2和NS2后，通过RSV摩擦接种沉默植株，研究这两者在RSV侵染过程中的作用；第三部分，利用VIGS技术沉默与OsSGS3同源的本氏烟NbSGS3，通过RSV摩擦接种沉默植株，检视SGS3对RSV是否具有抗病性，并根据研究结果，进一步构建OsSGS3干扰的转基因水稻。获得如下结果：
1发现NS2的亚细胞定位存在周期性变化，先定位在细胞核后运动到细胞核外；明确了NS2在细胞核内的具体定位及其与三种寄主的核仁和cajal body蛋白的互作情况。通过共定位观察发现NS2可以定位在柯浩体和核仁上，BiFC结果显示NS2可以与三种寄主编码的3个核仁和柯浩体指示蛋白即Atcoilin、NbFib2和OsFib2互作。
2明确了NS2和NbFib2在RSV侵染中的作用。RSV分别接种沉默了NS2和NbFib2的本氏烟，在沉默植株的接种叶中可以检测到病毒，而在非接种叶中检测不到病毒；病毒在沉默植株接种叶的积累量比对照即VIGS (空)植株低，这说明这两个基因影响了病毒的复制及系统侵染。在NbFib2沉默的本氏烟中，NS2早期的亚细胞定位发生了改变，与对照即健康植株或VIGS (空)接种植株相比，NS2形成的小颗粒不仅定位在细胞核，还分布在其它细胞器中。可见，RSV在进行系统侵染需要NS2和Fib2的参与，而NS2借助Fib2进入细胞核可能是RSV侵染和复制所需要的。
3发现沉默了与OsSGS3同源的NbSGS3的本氏烟植株，经RSV接种后，在发病时间和发病程度上均表现出对RSV感病性，根据这一结果利用RNA干扰技术构建并获得OsSGS3干扰的转基因水稻T0代，为进一步研究OsSGS3基因的抗病调控功能奠定基础。
Rice stripe disease is one of the most significant rice viral diseases, endangeringrice production of China. This disease is caused by Rice stripe virus (RSV), a typicalmember of the genus Tenuivirus. With the improvement of molecular biotechnologyin recent years, researches on RSV have been gradually advanced. Especially, afterdiscovery that two model plants, Nicotiana benthamiana (N. benthamiana) andArabidopsis thaliana (A. thaliana) can be infected by RSV, some difficulties andobstacles of researches in this field, such as long time requirment for growing rice andconstructing transgenic plants, were overcome. It has been reported that NS2, encodedby RSV, is a weak gene silencing suppressor, localizing in nucleus and interactingwith OsSGS3. SGS3, a type of relatively conservative gene, is a gene silencingsuppressor encoded by plant itself, and SGS3mutant A.thaliana is more susceptible tovirus. However, the functional mechanism of NS2during RSV infection on host plantsand the role of OsSGS3in RSV resistance are still unclear. In this PhD research, N.benthamiana and virus inducing gene silencing (VIGS) method were used to analyzethe pathogenicity functions of NS2and disease resistance conferred by OsSGS3toRSV; OsSGS3-RNAi transgenic plants were also constructed, laying the foundationfor further dissection of role of OsSGS3in rice resistance to RSV.
This dissertation comprises three parts of researches. In the first part,co-localization and BiFC analyses were conducted to verify the interaction of NS2with cajal body and nucleolus proteins in three different hosts, A. thaliana, N.benthamiana and rice. In the second part, the NbFib2and NS2genes were silenced byVIGS, then the silenced plants were infected with RSV, thereby the function of thesetwo genes in RSV infection was analyzed. In the last part, the function of SGS3toRSV was analyzed by silencing NbSGS3, a homolog of OsSGS3, in N. benthamianaand challenging the silenced plants with wild type RSV using the same approache asthe second part; OsSGS3-RNAi transgenic rice was constructed for further studies.Following is the summary of the research:
1. The subcellular localization of NS2varied over time.The protein first localizesin the nucleus and then migrates outside the nucleus. The subcellular localization ofNS2and the interaction between NS2and the cajal body and nuclelous proteins fromthree different hosts were identified. The co-localization results showed that NS2 localized in the cajal body and nucleolus. The BiFC assay revealed that NS2interacted with Atcoilin, NbFib2and OsFib2from three hosts.
2. The roles of NS2and NbFib2in RSV infection were identified. RSV can bedetected in the leaves of NS2-silenced plants or NbFib2-silenced plants inoculatedwith the virus, but not in uninoculated leaves of the same silenced plants. Theaccumulation of RSV in inoculated leaves of silenced plants was less than in theun-silenced plants (CK) that were the inoculated only with Agrobacterium containingempty VIGS vector, suggesting that NS2and NbFib2play important roles in RSVreplication and infection. NbFib2-silenced also altered the subcellular localization ofNS2in the earlier stage. Granules induced by NS2localized not only in nucleus, butalso in other organelles. These results indicate that both NS2and NbFib2are involvedin RSV systemic infection, NS2migrates into nucleus with assistance to completeRSV replication and systemic infection.
3. In comparison with the CK plants, the NbSGS3-silenced plants were moresusceptible to RSV infection as indicated by both latent period and diseased severity.Based on this result, OsSGS3-RNAi transgenic T0rice was constructed, laying afoundation for further characterization of the OsSGS3-conferred disease-resistance.
本文研究内容主要分为三部分。第一部分，通过共定位方法进一步确定NS2在细胞核内的具体位置，并通过双荧光互补实验(Bimolecular fluorescencecomplementation, BiFC)确定NS2与三种寄主即拟南芥、本氏烟和水稻的核仁和柯浩体的互作情况；第二部分，利用VIGS技术分别沉默基因NbFib2和NS2后，通过RSV摩擦接种沉默植株，研究这两者在RSV侵染过程中的作用；第三部分，利用VIGS技术沉默与OsSGS3同源的本氏烟NbSGS3，通过RSV摩擦接种沉默植株，检视SGS3对RSV是否具有抗病性，并根据研究结果，进一步构建OsSGS3干扰的转基因水稻。获得如下结果：
1发现NS2的亚细胞定位存在周期性变化，先定位在细胞核后运动到细胞核外；明确了NS2在细胞核内的具体定位及其与三种寄主的核仁和cajal body蛋白的互作情况。通过共定位观察发现NS2可以定位在柯浩体和核仁上，BiFC结果显示NS2可以与三种寄主编码的3个核仁和柯浩体指示蛋白即Atcoilin、NbFib2和OsFib2互作。
2明确了NS2和NbFib2在RSV侵染中的作用。RSV分别接种沉默了NS2和NbFib2的本氏烟，在沉默植株的接种叶中可以检测到病毒，而在非接种叶中检测不到病毒；病毒在沉默植株接种叶的积累量比对照即VIGS (空)植株低，这说明这两个基因影响了病毒的复制及系统侵染。在NbFib2沉默的本氏烟中，NS2早期的亚细胞定位发生了改变，与对照即健康植株或VIGS (空)接种植株相比，NS2形成的小颗粒不仅定位在细胞核，还分布在其它细胞器中。可见，RSV在进行系统侵染需要NS2和Fib2的参与，而NS2借助Fib2进入细胞核可能是RSV侵染和复制所需要的。
3发现沉默了与OsSGS3同源的NbSGS3的本氏烟植株，经RSV接种后，在发病时间和发病程度上均表现出对RSV感病性，根据这一结果利用RNA干扰技术构建并获得OsSGS3干扰的转基因水稻T0代，为进一步研究OsSGS3基因的抗病调控功能奠定基础。
Rice stripe disease is one of the most significant rice viral diseases, endangeringrice production of China. This disease is caused by Rice stripe virus (RSV), a typicalmember of the genus Tenuivirus. With the improvement of molecular biotechnologyin recent years, researches on RSV have been gradually advanced. Especially, afterdiscovery that two model plants, Nicotiana benthamiana (N. benthamiana) andArabidopsis thaliana (A. thaliana) can be infected by RSV, some difficulties andobstacles of researches in this field, such as long time requirment for growing rice andconstructing transgenic plants, were overcome. It has been reported that NS2, encodedby RSV, is a weak gene silencing suppressor, localizing in nucleus and interactingwith OsSGS3. SGS3, a type of relatively conservative gene, is a gene silencingsuppressor encoded by plant itself, and SGS3mutant A.thaliana is more susceptible tovirus. However, the functional mechanism of NS2during RSV infection on host plantsand the role of OsSGS3in RSV resistance are still unclear. In this PhD research, N.benthamiana and virus inducing gene silencing (VIGS) method were used to analyzethe pathogenicity functions of NS2and disease resistance conferred by OsSGS3toRSV; OsSGS3-RNAi transgenic plants were also constructed, laying the foundationfor further dissection of role of OsSGS3in rice resistance to RSV.
This dissertation comprises three parts of researches. In the first part,co-localization and BiFC analyses were conducted to verify the interaction of NS2with cajal body and nucleolus proteins in three different hosts, A. thaliana, N.benthamiana and rice. In the second part, the NbFib2and NS2genes were silenced byVIGS, then the silenced plants were infected with RSV, thereby the function of thesetwo genes in RSV infection was analyzed. In the last part, the function of SGS3toRSV was analyzed by silencing NbSGS3, a homolog of OsSGS3, in N. benthamianaand challenging the silenced plants with wild type RSV using the same approache asthe second part; OsSGS3-RNAi transgenic rice was constructed for further studies.Following is the summary of the research:
1. The subcellular localization of NS2varied over time.The protein first localizesin the nucleus and then migrates outside the nucleus. The subcellular localization ofNS2and the interaction between NS2and the cajal body and nuclelous proteins fromthree different hosts were identified. The co-localization results showed that NS2 localized in the cajal body and nucleolus. The BiFC assay revealed that NS2interacted with Atcoilin, NbFib2and OsFib2from three hosts.
2. The roles of NS2and NbFib2in RSV infection were identified. RSV can bedetected in the leaves of NS2-silenced plants or NbFib2-silenced plants inoculatedwith the virus, but not in uninoculated leaves of the same silenced plants. Theaccumulation of RSV in inoculated leaves of silenced plants was less than in theun-silenced plants (CK) that were the inoculated only with Agrobacterium containingempty VIGS vector, suggesting that NS2and NbFib2play important roles in RSVreplication and infection. NbFib2-silenced also altered the subcellular localization ofNS2in the earlier stage. Granules induced by NS2localized not only in nucleus, butalso in other organelles. These results indicate that both NS2and NbFib2are involvedin RSV systemic infection, NS2migrates into nucleus with assistance to completeRSV replication and systemic infection.
3. In comparison with the CK plants, the NbSGS3-silenced plants were moresusceptible to RSV infection as indicated by both latent period and diseased severity.Based on this result, OsSGS3-RNAi transgenic T0rice was constructed, laying afoundation for further characterization of the OsSGS3-conferred disease-resistance.
引文
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