SRBSDV和RRSV在介体飞虱体内的侵染机理
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摘要
近年来由南方水稻黑条矮缩病毒（Southern rice black-streaked dwarf virus,SRBSDV）引起的南方水稻黑条矮缩病在我国南方稻区暴发流行，已成为我国目前水稻（Oryza sativa）生产上的重要病害之一。同时由水稻齿叶矮缩病毒（Riceragged stunt virus, RRSV）引起的水稻齿叶矮缩病也在我国南方稻区蔓延发生。SRBSDV是一种新发现的水稻病毒，属于呼肠孤病毒科（Reoviridae）斐济病毒属（Fijivirus）的一个暂定种，由新的介体白背飞虱（Sogatella furcifera）以持久增殖型方式传播。RRSV是一种再次发生的老病毒，是呼肠孤病毒科（Reoviridae）水稻病毒属（Oryzavirus）的代表种，由介体褐飞虱（Nilaparvata lugens）以持久增殖型方式传播。目前，国内的科研工作者已对这两种水稻病毒的介体传毒特性做了大量的研究，然而关于病毒在介体飞虱体内的侵染机理知之甚少，有关病毒在介体体内的侵染循回过程以及复制和扩散机理还未曾报道。
本研究首先通过免疫荧光标记技术系统研究了SRBSDV在其介体昆虫体内的侵染循回过程。明确了SRBSDV在白背飞虱取食过程中首先进入中肠肠腔；随后少量的病毒粒体侵入中肠上皮细胞并进行复制；增殖后的病毒从中肠上皮细胞向中肠表面的肌肉组织扩散；病毒在中肠表面沿着环肌和纵肌向邻近的器官扩散；饲毒后6d，部分白背飞虱体内SRBSDV已通过血淋巴扩散到唾液腺；饲毒后8d，病毒在白背飞虱唾液腺中大量增殖，达到了系统性侵染，从而完成了整个侵染循回过程。SRBSDV与白背飞虱和灰飞虱的亲和性具有明显的差别，这可能与病毒在两种介体体内的侵染循回过程不同有关。通过共聚焦显微镜观察SRBSDV在饲毒后25d的灰飞虱消化系统的分布，发现灰飞虱的中肠内可以检测到病毒，但在唾液腺中检测不到病毒，明确了SRBSDV受到中肠释放屏障的阻碍无法扩散到唾液腺，从而不能被灰飞虱有效传播。
SRBSDV在介体昆虫内的侵染会诱导形成提供病毒复制和装配场所的球状或纤维状的电子致密内含体——病毒原质（viroplasm）。为进一步研究SRBSDV在介体昆虫体内的侵染和增殖机制，本研究在SRBSDV侵染的白背飞虱培养细胞中观察到病毒编码的非结构蛋白P9-1分布在病毒原质基质中。子代病毒粒体和病毒复制产生的病毒RNA积累在P9-1分布的病毒原质基质中。在体外体系中，SRBSDV P9-1可独立形成类似于病毒原质基质的内含体。这些结果表明，SRBSDV P9-1是形成病毒原质基质的必需组分，形成的病毒原质是SRBSDV进行病毒RNA复制和病毒粒体装配的场所。为了进一步验证SRBSDV P9-1在病毒侵染介体过程中的功能，本研究首次将人工合成dsRNA诱导的RNAi技术与白背飞虱培养细胞技术相结合研究P9-1的功能。通过转染针对P9-1基因的人工合成的dsRNA干扰P9-1基因在介体培养细胞中的表达显著抑制病毒原质的形成和病毒在细胞中的侵染。将人工合成的P9-1基因dsRNA通过膜饲喂法饲喂白背飞虱，抑制了SRBSDV在白背飞虱体内的侵染和复制，从而阻碍了白背飞虱传播病毒。因此，SRBSDV P9-1是病毒在介体昆虫体内有效侵染和复制所必需的。
SRBSDV侵染白背飞虱后必须从初侵染的中肠上皮细胞扩散到唾液腺才能被介体有效传播，因此病毒在介体内的扩散也是其完成侵染循回的关键环节。本研究在SRBSDV侵染的白背飞虱培养细胞中观察到由非结构蛋白P7-1形成的包裹着病毒粒体的管状结构可伸出侵染细胞的表面。在体外实验中，单独表达的P7-1能在Sf9细胞中形成伸出细胞的小管结构，推测其参与病毒在介体昆虫体内的扩散。为了验证P7-1蛋白在病毒扩散过程中的作用，本研究通过病毒抗体中和培养细胞中游离病毒粒体后，仍可观察到P7-1管状结构从初次侵染的细胞延伸到邻近的健康细胞。体外转染人工合成的针对P7-1基因的dsRNA有效抑制P7-1管状结构的形成，阻碍SRBSDV在细胞间的扩散，表明P7-1管状结构是SRBSDV在白背飞虱培养细胞间安全扩散的重要通道。
P7-1管状结构是否介导病毒在白背飞虱体内扩散？通过免疫荧光标记分析发现P7-1管状结构在白背飞虱体内的扩散途径与病毒的侵染循回过程一致，P7-1管状结构从白背飞虱中肠上皮细胞首先扩散到中肠表面的环肌，并沿着环肌横向扩散，当遇到与环肌交叉的纵肌后开始沿着纵肌纵向扩散，最后扩散到整个中肠和邻近器官的表面。在P7-1管状结构到达的部位病毒才能进行复制，病毒的复制滞后于P7-1管状结构的扩散。通过酵母双杂交系统和免疫共沉淀技术发现P7-1与肌动蛋白（Actin）存在特异性互作。因此，P7-1管状结构具备了在消化道外层肌纤维上运动的能力。通过膜饲喂法饲喂人工合成的针对P7-1基因的dsRNA，有效抑制P7-1管状结构的形成，阻碍病毒在白背飞虱体内的扩散，最终阻碍了白背飞虱传播SRBSDV的能力，表明SRBSDV P7-1是病毒在白背飞虱体内有效扩散所必需的。
本研究对RRSV在其介体褐飞虱体内的侵染循回过程和复制机理进行了分析。首先通过免疫荧光标记技术明确了RRSV在褐飞虱体内的侵染循回途径与SRBSDV在白背飞虱体内的侵染循回途径大体相似。在病毒侵染褐飞虱过程中，由非结构蛋白Pns10诱导形成的病毒原质在中肠上皮细胞、中肠外层肌肉组织和唾液腺等组织均有分布。通过膜饲喂体外合成的针对Pns10基因的dsRNA抑制了病毒原质在介体昆虫体内的形成，阻碍病毒在褐飞虱体内的侵染和复制，证明了Pns10蛋白是RRSV侵染介体褐飞虱所必需的。
综上所述，本研究首次阐明了SRBSDV和RRSV在不同飞虱体内的侵染循回过程；首次将RNAi技术与介体昆虫培养细胞相结合研究植物病毒编码蛋白的功能，明确了SRBSDV非结构蛋白P9-1和RRSV非结构蛋白Pns10是形成病毒原质基质的组分，参与病毒在介体内的复制，SRBSDV非结构蛋白P7-1蛋白是形成包裹病毒粒体的管状结构的组分，参与病毒在介体内的扩散。初步解析了SRBSDV和RRSV适应于相应介体昆虫高效传毒的分子机制，为利用RNAi原理培育转基因水稻防治病毒病的发生提供了靶标基因及理论依据。
Southern rice black-streaked dwarf disease caused by southern rice black-streakeddwarf virus (SRBSDV) has recently prevailed in southern China and become one ofthe most damaging viral diseases of rice. SRBSDV is a newly proposed Fijivirusspecies in the family Reoviridae, which was transmitted mainly by white backedplanthopper (Sogatella furcifera, WBPH） in a persistent propagative manner.Meanwhile, Rice ragged stunt disease caused by rice ragged stunt virus (RRSV) hasrecently spread in southern China. RRSV is a well-established Oryzavirus species inthe family Reoviridae. It is transmitted by brown planthopper (Nilaparvata lugens,BPH), also in a persistent propagative manner. In recent years, transmissioncharacteristics of the two rice viruses by planthoppers have been studied by manyresearchers in China. However, few studies have been conducted on the mechanismsby which the two viruses infect their vector planthoppers. The infection route,replication and spread of viruses in insect vectors are still poorly understood.
In this study, we investigated the infection route of SRBSDV in the digestivesystem of WBPH by immunofluorescence. SRBSDV particles initially traveledthrough the oesophagus into the midgut lumen of WBPH after insect acquisition ofthe virus from SRBSDV-infected rice plants. Then, only a few virus particles infectedand replicated in the epithelial cells of the midgut. At4days post-first access todiseased plants (padp), SRBSDV traversed the basal lamina and infected the visceralmuscle tissues encircling the midgut epithelium. From then on, SRBSDV spreadalone the circular and longitudinal muscles of the midgut to other organs. At6dayspadp, SRBSDV spread to salivary glands in some insects. At8days padp, SRBSDVinfected the whole alimentary canal and replicated in the salivary glands of WBPH.The compatibility between SRBSDV and WBPH is significantly different from thatbetween SRBSDV and SBPH. It may be related to the different infection route ofSRBSDV in different vector insects. The localization of SRBSDV in digestive systemof SBPHs at25days padp was detected by immunofluorescence microscopy. Wecould observe SRBSDV within the midgut, but not in the salivary glands of SBPHs.This result suggested that SRBSDV was restricted in midgut and failed to spread intothe salivary glands because of the midgut dissemination barrier in SBPHs, which ledto the SBPH failed to transmit SRBSDV to rice.
During the infection of SRBSDV, viroplasms, the punctate inclusions for virusreplication and assembly of progeny virions, were formed in the alimentary canal andsalivary glands of BPHs. To study the infection mechanism of SRBSDV, we usedimmunofluorescence microscopy to observe the localization of P9-1in virus-infectedVCMs of WBPHs. P9-1formed punctate inclusions in VCMs, and progeny viralparticles and viral mRNA accumulated within the viroplasm matrix of P9-1. P9-1alsocould form viroplasm-like inclusions in the Sf9cells. All these results suggested thatP9-1is a constituent of the matrix of viroplasm and viral inclusions formed by P9-1were the sites for synthesising viral RNA and assembling viral particles. To furtherconfirm the function of P9-1in viral replication, the RNA interference induced bysynthesized dsRNA was used to investigate gene functions of P9-1in insect culturecells. By knocking down the expression of P9-1with synthesized dsRNA of P9-1, theformation of viroplasm and viral infection were significantly inhibited in the VCMs.Meanwhile, ingestion of dsRNAs from the P9-1gene via membrane feeding alsostrongly inhibited viral infection and replication in intact WBPHs. These resultssuggested that P9-1was essential for the infection and replication of SRBSDV inWBPHs.
SRBSDV can only be effectively transmitted when it spread from the midgutepithelial cells, the primary site of virus entry into insect cells, to salivary glands. Thespread of SRBSDV in the insect vector plays a crucial role in the infection route. Inthis study, we observed that the tubular structures formed by non-structure proteinP7-1contained viral particles and scattered the surface of infected cells or extendedtoward neighboring cells. P7-1of SRBSDV also has the ability to form tubulesgrowing from the cell surface in the Sf9cells. Based on these results, we conjecturedthat the P7-1was involved in the cell-to-cell transmission of SRBSDV. To determinethe function of P7-1related to the spread of SRBSDV, we blocked the virus that hadbeen released into or was present in VCMs with virus-neutralizing antibodies. TheP7-1tubular structures were also extended from the primary cell toward neighboringhealthy cells. When the expression of P7-1was knocked down by RNAi induced bydsRNA specific for P7-1gene, the formation of tubules was inhibited and SRBSDVfailed to spread to the neighboring cells. All these results suggested that the P7-1participated in the cell-to-cell spread of SRBSDV in cells of WBPHs.
Whether the P7-1participates in the spread of SRBSDV in WBPHs? We detectedthe localization of P7-1in WBPHs by confocal microscopy. The distribution oftubular structures formed by P7-1is correlated with the infection route of SRBSDV. P7-1firstly appeared in epithelial cells, then spread from the epithelial cells to circularmuscle of midgut and moved alone it. Tubules subsequently spread to the longitudinalmuscle that intersected with circular muscle and disseminated from the midgut toother organs. Virus started to replicate on the sites where tubules had arrived before.We confirmed that the P7-1specifically interacted with Actin of WBPH by yeast twohybrid experiments and co-immunoprecipitation, suggesting that the tubule formed byP7-1utilized the interaction with Actin of WBPH to spread on muscle of midgut.When the expression of P7-1was strongly inhibited by feeding synthesized dsRNA ofP7-1via membrane feeding, the formation of P7-1tubular structure was inhibited.This prevented the spread of SRBSDV in the digestive system of WBPHs, whichresulted in the fact that the tested WBPH failed to transmit virus to rice. All theseresults suggested that the P7-1was essential for the spread of SRBSDV in WBPH.
We also determined the infection route and the replication mechanism of RRSV ininfected BPHs. We firstly analyzed the infection route of RRSV in BPHs byimmunofluorescence and found that it was roughly similar with that of SRBSDV.During the infection of RRSV, non-structure protein Pns10formed viroplasm inmidgut epithelial, muscle and salivary glands of RRSV-infected BPHs. To furtherconfirm the function of Pns10in viral infection, Second-instar nymphs of BPHs werefed synthesized dsRNA of Pns10by membrane feeding before acquisition ofSRBSDV. As a result, RNAi targeting Pns10significantly inhibited the formation ofviroplasm and prevented viral infection and replication in intact BPHs. These resultsindicate that Pns10is essential for infection of RRSV in BPHs.
In all, the infection routes of SRBSDV and RRSV in different vector planthopperswere elucidated for the first time in this study. It was the first time to use RNAistrategy to investigate gene functions of plant viruses in insect culture cells. UsingRNAi induced by synthesized dsRNA, we found that P9-1of SRBSDV and Pns10ofRRSV were the constituent of the viroplasm matrix and were essential for viralreplication. Tubular structures formed by P7-1of SRBSDV contained viral particlesand participated in viral spread in insect. These studies are useful for understandingthe efficiently transmission mechanism of SRBSDV and RRSV in their vectors, andprovided theoretical basis and new targets to control viral diseases of rice by RNAistrategy.
本研究首先通过免疫荧光标记技术系统研究了SRBSDV在其介体昆虫体内的侵染循回过程。明确了SRBSDV在白背飞虱取食过程中首先进入中肠肠腔；随后少量的病毒粒体侵入中肠上皮细胞并进行复制；增殖后的病毒从中肠上皮细胞向中肠表面的肌肉组织扩散；病毒在中肠表面沿着环肌和纵肌向邻近的器官扩散；饲毒后6d，部分白背飞虱体内SRBSDV已通过血淋巴扩散到唾液腺；饲毒后8d，病毒在白背飞虱唾液腺中大量增殖，达到了系统性侵染，从而完成了整个侵染循回过程。SRBSDV与白背飞虱和灰飞虱的亲和性具有明显的差别，这可能与病毒在两种介体体内的侵染循回过程不同有关。通过共聚焦显微镜观察SRBSDV在饲毒后25d的灰飞虱消化系统的分布，发现灰飞虱的中肠内可以检测到病毒，但在唾液腺中检测不到病毒，明确了SRBSDV受到中肠释放屏障的阻碍无法扩散到唾液腺，从而不能被灰飞虱有效传播。
SRBSDV在介体昆虫内的侵染会诱导形成提供病毒复制和装配场所的球状或纤维状的电子致密内含体——病毒原质（viroplasm）。为进一步研究SRBSDV在介体昆虫体内的侵染和增殖机制，本研究在SRBSDV侵染的白背飞虱培养细胞中观察到病毒编码的非结构蛋白P9-1分布在病毒原质基质中。子代病毒粒体和病毒复制产生的病毒RNA积累在P9-1分布的病毒原质基质中。在体外体系中，SRBSDV P9-1可独立形成类似于病毒原质基质的内含体。这些结果表明，SRBSDV P9-1是形成病毒原质基质的必需组分，形成的病毒原质是SRBSDV进行病毒RNA复制和病毒粒体装配的场所。为了进一步验证SRBSDV P9-1在病毒侵染介体过程中的功能，本研究首次将人工合成dsRNA诱导的RNAi技术与白背飞虱培养细胞技术相结合研究P9-1的功能。通过转染针对P9-1基因的人工合成的dsRNA干扰P9-1基因在介体培养细胞中的表达显著抑制病毒原质的形成和病毒在细胞中的侵染。将人工合成的P9-1基因dsRNA通过膜饲喂法饲喂白背飞虱，抑制了SRBSDV在白背飞虱体内的侵染和复制，从而阻碍了白背飞虱传播病毒。因此，SRBSDV P9-1是病毒在介体昆虫体内有效侵染和复制所必需的。
SRBSDV侵染白背飞虱后必须从初侵染的中肠上皮细胞扩散到唾液腺才能被介体有效传播，因此病毒在介体内的扩散也是其完成侵染循回的关键环节。本研究在SRBSDV侵染的白背飞虱培养细胞中观察到由非结构蛋白P7-1形成的包裹着病毒粒体的管状结构可伸出侵染细胞的表面。在体外实验中，单独表达的P7-1能在Sf9细胞中形成伸出细胞的小管结构，推测其参与病毒在介体昆虫体内的扩散。为了验证P7-1蛋白在病毒扩散过程中的作用，本研究通过病毒抗体中和培养细胞中游离病毒粒体后，仍可观察到P7-1管状结构从初次侵染的细胞延伸到邻近的健康细胞。体外转染人工合成的针对P7-1基因的dsRNA有效抑制P7-1管状结构的形成，阻碍SRBSDV在细胞间的扩散，表明P7-1管状结构是SRBSDV在白背飞虱培养细胞间安全扩散的重要通道。
P7-1管状结构是否介导病毒在白背飞虱体内扩散？通过免疫荧光标记分析发现P7-1管状结构在白背飞虱体内的扩散途径与病毒的侵染循回过程一致，P7-1管状结构从白背飞虱中肠上皮细胞首先扩散到中肠表面的环肌，并沿着环肌横向扩散，当遇到与环肌交叉的纵肌后开始沿着纵肌纵向扩散，最后扩散到整个中肠和邻近器官的表面。在P7-1管状结构到达的部位病毒才能进行复制，病毒的复制滞后于P7-1管状结构的扩散。通过酵母双杂交系统和免疫共沉淀技术发现P7-1与肌动蛋白（Actin）存在特异性互作。因此，P7-1管状结构具备了在消化道外层肌纤维上运动的能力。通过膜饲喂法饲喂人工合成的针对P7-1基因的dsRNA，有效抑制P7-1管状结构的形成，阻碍病毒在白背飞虱体内的扩散，最终阻碍了白背飞虱传播SRBSDV的能力，表明SRBSDV P7-1是病毒在白背飞虱体内有效扩散所必需的。
本研究对RRSV在其介体褐飞虱体内的侵染循回过程和复制机理进行了分析。首先通过免疫荧光标记技术明确了RRSV在褐飞虱体内的侵染循回途径与SRBSDV在白背飞虱体内的侵染循回途径大体相似。在病毒侵染褐飞虱过程中，由非结构蛋白Pns10诱导形成的病毒原质在中肠上皮细胞、中肠外层肌肉组织和唾液腺等组织均有分布。通过膜饲喂体外合成的针对Pns10基因的dsRNA抑制了病毒原质在介体昆虫体内的形成，阻碍病毒在褐飞虱体内的侵染和复制，证明了Pns10蛋白是RRSV侵染介体褐飞虱所必需的。
综上所述，本研究首次阐明了SRBSDV和RRSV在不同飞虱体内的侵染循回过程；首次将RNAi技术与介体昆虫培养细胞相结合研究植物病毒编码蛋白的功能，明确了SRBSDV非结构蛋白P9-1和RRSV非结构蛋白Pns10是形成病毒原质基质的组分，参与病毒在介体内的复制，SRBSDV非结构蛋白P7-1蛋白是形成包裹病毒粒体的管状结构的组分，参与病毒在介体内的扩散。初步解析了SRBSDV和RRSV适应于相应介体昆虫高效传毒的分子机制，为利用RNAi原理培育转基因水稻防治病毒病的发生提供了靶标基因及理论依据。
Southern rice black-streaked dwarf disease caused by southern rice black-streakeddwarf virus (SRBSDV) has recently prevailed in southern China and become one ofthe most damaging viral diseases of rice. SRBSDV is a newly proposed Fijivirusspecies in the family Reoviridae, which was transmitted mainly by white backedplanthopper (Sogatella furcifera, WBPH） in a persistent propagative manner.Meanwhile, Rice ragged stunt disease caused by rice ragged stunt virus (RRSV) hasrecently spread in southern China. RRSV is a well-established Oryzavirus species inthe family Reoviridae. It is transmitted by brown planthopper (Nilaparvata lugens,BPH), also in a persistent propagative manner. In recent years, transmissioncharacteristics of the two rice viruses by planthoppers have been studied by manyresearchers in China. However, few studies have been conducted on the mechanismsby which the two viruses infect their vector planthoppers. The infection route,replication and spread of viruses in insect vectors are still poorly understood.
In this study, we investigated the infection route of SRBSDV in the digestivesystem of WBPH by immunofluorescence. SRBSDV particles initially traveledthrough the oesophagus into the midgut lumen of WBPH after insect acquisition ofthe virus from SRBSDV-infected rice plants. Then, only a few virus particles infectedand replicated in the epithelial cells of the midgut. At4days post-first access todiseased plants (padp), SRBSDV traversed the basal lamina and infected the visceralmuscle tissues encircling the midgut epithelium. From then on, SRBSDV spreadalone the circular and longitudinal muscles of the midgut to other organs. At6dayspadp, SRBSDV spread to salivary glands in some insects. At8days padp, SRBSDVinfected the whole alimentary canal and replicated in the salivary glands of WBPH.The compatibility between SRBSDV and WBPH is significantly different from thatbetween SRBSDV and SBPH. It may be related to the different infection route ofSRBSDV in different vector insects. The localization of SRBSDV in digestive systemof SBPHs at25days padp was detected by immunofluorescence microscopy. Wecould observe SRBSDV within the midgut, but not in the salivary glands of SBPHs.This result suggested that SRBSDV was restricted in midgut and failed to spread intothe salivary glands because of the midgut dissemination barrier in SBPHs, which ledto the SBPH failed to transmit SRBSDV to rice.
During the infection of SRBSDV, viroplasms, the punctate inclusions for virusreplication and assembly of progeny virions, were formed in the alimentary canal andsalivary glands of BPHs. To study the infection mechanism of SRBSDV, we usedimmunofluorescence microscopy to observe the localization of P9-1in virus-infectedVCMs of WBPHs. P9-1formed punctate inclusions in VCMs, and progeny viralparticles and viral mRNA accumulated within the viroplasm matrix of P9-1. P9-1alsocould form viroplasm-like inclusions in the Sf9cells. All these results suggested thatP9-1is a constituent of the matrix of viroplasm and viral inclusions formed by P9-1were the sites for synthesising viral RNA and assembling viral particles. To furtherconfirm the function of P9-1in viral replication, the RNA interference induced bysynthesized dsRNA was used to investigate gene functions of P9-1in insect culturecells. By knocking down the expression of P9-1with synthesized dsRNA of P9-1, theformation of viroplasm and viral infection were significantly inhibited in the VCMs.Meanwhile, ingestion of dsRNAs from the P9-1gene via membrane feeding alsostrongly inhibited viral infection and replication in intact WBPHs. These resultssuggested that P9-1was essential for the infection and replication of SRBSDV inWBPHs.
SRBSDV can only be effectively transmitted when it spread from the midgutepithelial cells, the primary site of virus entry into insect cells, to salivary glands. Thespread of SRBSDV in the insect vector plays a crucial role in the infection route. Inthis study, we observed that the tubular structures formed by non-structure proteinP7-1contained viral particles and scattered the surface of infected cells or extendedtoward neighboring cells. P7-1of SRBSDV also has the ability to form tubulesgrowing from the cell surface in the Sf9cells. Based on these results, we conjecturedthat the P7-1was involved in the cell-to-cell transmission of SRBSDV. To determinethe function of P7-1related to the spread of SRBSDV, we blocked the virus that hadbeen released into or was present in VCMs with virus-neutralizing antibodies. TheP7-1tubular structures were also extended from the primary cell toward neighboringhealthy cells. When the expression of P7-1was knocked down by RNAi induced bydsRNA specific for P7-1gene, the formation of tubules was inhibited and SRBSDVfailed to spread to the neighboring cells. All these results suggested that the P7-1participated in the cell-to-cell spread of SRBSDV in cells of WBPHs.
Whether the P7-1participates in the spread of SRBSDV in WBPHs? We detectedthe localization of P7-1in WBPHs by confocal microscopy. The distribution oftubular structures formed by P7-1is correlated with the infection route of SRBSDV. P7-1firstly appeared in epithelial cells, then spread from the epithelial cells to circularmuscle of midgut and moved alone it. Tubules subsequently spread to the longitudinalmuscle that intersected with circular muscle and disseminated from the midgut toother organs. Virus started to replicate on the sites where tubules had arrived before.We confirmed that the P7-1specifically interacted with Actin of WBPH by yeast twohybrid experiments and co-immunoprecipitation, suggesting that the tubule formed byP7-1utilized the interaction with Actin of WBPH to spread on muscle of midgut.When the expression of P7-1was strongly inhibited by feeding synthesized dsRNA ofP7-1via membrane feeding, the formation of P7-1tubular structure was inhibited.This prevented the spread of SRBSDV in the digestive system of WBPHs, whichresulted in the fact that the tested WBPH failed to transmit virus to rice. All theseresults suggested that the P7-1was essential for the spread of SRBSDV in WBPH.
We also determined the infection route and the replication mechanism of RRSV ininfected BPHs. We firstly analyzed the infection route of RRSV in BPHs byimmunofluorescence and found that it was roughly similar with that of SRBSDV.During the infection of RRSV, non-structure protein Pns10formed viroplasm inmidgut epithelial, muscle and salivary glands of RRSV-infected BPHs. To furtherconfirm the function of Pns10in viral infection, Second-instar nymphs of BPHs werefed synthesized dsRNA of Pns10by membrane feeding before acquisition ofSRBSDV. As a result, RNAi targeting Pns10significantly inhibited the formation ofviroplasm and prevented viral infection and replication in intact BPHs. These resultsindicate that Pns10is essential for infection of RRSV in BPHs.
In all, the infection routes of SRBSDV and RRSV in different vector planthopperswere elucidated for the first time in this study. It was the first time to use RNAistrategy to investigate gene functions of plant viruses in insect culture cells. UsingRNAi induced by synthesized dsRNA, we found that P9-1of SRBSDV and Pns10ofRRSV were the constituent of the viroplasm matrix and were essential for viralreplication. Tubular structures formed by P7-1of SRBSDV contained viral particlesand participated in viral spread in insect. These studies are useful for understandingthe efficiently transmission mechanism of SRBSDV and RRSV in their vectors, andprovided theoretical basis and new targets to control viral diseases of rice by RNAistrategy.
引文
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