口蹄疫病毒3C（D）蛋白与宿主细胞因子Fascin-1相互作用研究
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摘要
口蹄疫病毒(FMDV)是小RNA病毒科口蹄疫病毒属的模式毒株,有七种不同血清型：A、O、C、Asia1、SAT1、SAT2和SAT3,以及进化成许多亚型。它的基因组是一条单股正链RNA,长度超过8kb,其5’端共价结合一个23-24氨基酸的基因组连接蛋白3B。FMDV基因组的复制方式同其他小RNA病毒科的病毒一样利用一条与正链互补的负链RNA作为其复制中间体。为有效完成其生命周期,不仅需要一些病毒蛋白,也需要一些宿主因子在病毒感染的大部分阶段发挥重要作用,因而鉴定这些宿主因子和它们的功能长期以来都被作为一个重要的研究目标。
一、口蹄疫病毒全长cDNA克隆的构建
为了保持口蹄疫病毒的基因组稳定性和保存遗传信息,我们成功地构建了O型口蹄疫全长cDNA克隆质粒pBluescript-FMDV.依据FMDV基因组本身带有的三个内切酶位点分成4个片段分别克隆后整合成完整的全长cDNA克隆,在其3’端引入25个poly(A)尾,并利用常规RT-PCR获得了15个poly(C)的5'UTR。
二、用于检测口蹄疫病毒正负链的双通道实时荧光定量RT-PCR的建立
为了更好地研究口蹄疫病毒的感染和复制,我们建立了一种简单而经济的双通道实时荧光定量RT-PCR技术,同时检测和定量病毒的正负链水平。引物和Taqman探针分别选自在亚型中最为保守的病毒基因2B和3D编码区。我们将此技术成功应用到了急性感染细胞、组织样品和单细胞样品中。在体外病毒感染早期,复制水平在感染后9h达到峰值而负链只有到感染后3h才能观察到；在FMDV感染的动物体内不同组织(肝脏、肾脏和脾脏)的正负链比值的动力学十分相似,反映出它们的复制动力学也相似；187个单细胞中有55个均检测出正链和负链,其比值从15.6到1463.4不等,展现出细胞个体之间的差异性,揭示出单细胞中复制活性有比较大的差别。这些证实了我们建立的双通道实时荧光定量RT-PCR有效而且可靠。
三、口蹄疫病毒3CDpro与BHK-21细胞因子Fascin-1的相互作用
口蹄疫病毒的复制和致病机制既与病毒编码的蛋白相关也涉及一些细胞因子。我们利用串联亲和层析纯化技术从BHK-21细胞中分离出可能与重组的前体蛋白△3CDpro(在其3C结构域C端缺失了30个氨基酸,因而丧失了其自我切割成熟的能力)相互作用的宿主蛋白,通过质谱分析出细胞蛋白Fascin-1。通过免疫共沉淀实验证实了△3CDpro及△3Cpro与Fascin在293T细胞内相互作用,但3Dpro却不能,进一步GSTpull-down实验也证实未缺失蛋白3CDpro和3CPro同样能在体外与Fascin蛋白直接相互作用。荧光共聚焦显微镜观察它们均定位于BHK-21细胞质中并且有很好的共定位。上调或下调Fascin蛋白表达能分别抑制或促进FMDV基因组复制,尤其是负链RNA的合成,而未对PV造成影响；FMDV感染早期(推测为复制起始时)能降低细胞内Fascin基因mRNA水平。
Foot-and-mouth disease (FMD) is one of the most highly contagious diseases of animals or humans, and the causative agent foot-and-mouth disease virus (FMDV) rapidly replicates and spreads within the infected animal, among in-contact susceptible animals, and by aerosol. FMDV is the prototype member of the Aphthovirus genus of the family Picornaviridae. The virus exists in the form of seven different serotypes:A, O, C, Asial, and South African Territories 1 (SAT1), SAT2, and SAT3, but a large number of subtypes have evolved within each serotype. Its genome is a single-stranded, positive-sense RNA which is over 8000 bases in length covalently bound at 5'-terminus to a 23-24 amino acid residue genome-linked protein, 3B. Replication of the FMDV genome RNA follows the same general pattern as replication of other members in the Picornaviridae family which occurs via a complementary negative strand RNA intermediate. To efficiently complete the life cycle of FMDV, not only viral proteins are involved, but also some host factors play important roles in most steps of viral infection, and identification of such host factors and their contributions has long been recognized as an important frontier.
1. Construction of FMDV full-length cDNA clone.
To stabilize FMDV genome and its genetical information, we successfully constructed type O FMDV full-length cDNA clone vector, pBluescript-FMDV. Basing on three incision enzyme sites within viral genome, four fragments were cloned into pBluescriptⅡKS(+) vector sequentially and then integrated which has artificial 25 poly(A) in 3'-terminal and 15 poly(C) within 5'UTR.
2. Establishment of a duplex quantitative real-time RT-PCR assay for simultaneous detection and quantitation of foot-and-mouth disease viral positive-stranded RNAs and negative-stranded RNAs.
Simplified, cost-effective, two-step duplex quantitative real-time RT-PCR assay was shown to detect and quantify foot-and-mouth disease virus positive-stranded RNAs and negative-stranded RNAs simultaneously for improved investigation of the state of virus infection and replication. The primers and Taqman probes were selected from the coding regions of 2B gene and 3D gene respectively, which have the least variations among serotypes. Cells infected acutely, tissue samples and single cell samples were used for evaluation of the assay. At the early stages of virus infection in vitro, the replication level reached a peak at 9 h.p.i. and the negative strands were detectable until 3 h.p.i. The kinetics of ratios of positive strands to negative strands (+RNA/-RNA) in vivo in the liver, kidney and spleen were similar, which demonstrated that the replication dynamics were similar in the three organs.55 single cell samples out of 187 were positive by both positive strands qPCR and negative strands qPCR, the ratios (+RNA/-RNA) ranged from 15.6 to 1463.4 which showed considerable difference among single cell samples, indicating that active viral replication differs greatly in single cells. A duplex quantitative real-time RT-PCR was validated as effective and reliable.
3. Interaction between FMDV 3CDpro and host factor Fascin-1 in BHK-21 cells.
Both virus-encoded proteins and cellular factors are involved in foot-and-mouth disease virus (FMDV) replication and pathogenesis. Tandem affinity purification was used in our study to purify cellular interactors of FMDV recombined 3 CD premature protein,△3CDpro (deletion of C-terminal 30 amino acids of 3C fragment in order to inactivate its auto-cleaving function), one of which was then identified as Fascin-1 by mass spectrum. The interaction between Fascin-1 and△3CDpro or△3Cpro rather than 3Dpro was conformed by co-immunoprecipitation assays. Results of confocal immunofluorescence analysis showed that they were well co-located in the cytoplasm of BHK-21 cells. GST pull down assay demonstrated further that the fascin directly interact with the 3CDpro as well as one of its mature protein 3Cpro in vitro. Overexpression or down-regulated expression of fascin-1 could inhibit or promote the replication of FMDV in BHK-21 rather than PV in Vero cells, especially synthesis of negative-strand RNAs. Furthermore, the levels of fascin mRNA decreased in early step of FMDV infection (maybe the initiation of replication).
一、口蹄疫病毒全长cDNA克隆的构建
为了保持口蹄疫病毒的基因组稳定性和保存遗传信息,我们成功地构建了O型口蹄疫全长cDNA克隆质粒pBluescript-FMDV.依据FMDV基因组本身带有的三个内切酶位点分成4个片段分别克隆后整合成完整的全长cDNA克隆,在其3’端引入25个poly(A)尾,并利用常规RT-PCR获得了15个poly(C)的5'UTR。
二、用于检测口蹄疫病毒正负链的双通道实时荧光定量RT-PCR的建立
为了更好地研究口蹄疫病毒的感染和复制,我们建立了一种简单而经济的双通道实时荧光定量RT-PCR技术,同时检测和定量病毒的正负链水平。引物和Taqman探针分别选自在亚型中最为保守的病毒基因2B和3D编码区。我们将此技术成功应用到了急性感染细胞、组织样品和单细胞样品中。在体外病毒感染早期,复制水平在感染后9h达到峰值而负链只有到感染后3h才能观察到；在FMDV感染的动物体内不同组织(肝脏、肾脏和脾脏)的正负链比值的动力学十分相似,反映出它们的复制动力学也相似；187个单细胞中有55个均检测出正链和负链,其比值从15.6到1463.4不等,展现出细胞个体之间的差异性,揭示出单细胞中复制活性有比较大的差别。这些证实了我们建立的双通道实时荧光定量RT-PCR有效而且可靠。
三、口蹄疫病毒3CDpro与BHK-21细胞因子Fascin-1的相互作用
口蹄疫病毒的复制和致病机制既与病毒编码的蛋白相关也涉及一些细胞因子。我们利用串联亲和层析纯化技术从BHK-21细胞中分离出可能与重组的前体蛋白△3CDpro(在其3C结构域C端缺失了30个氨基酸,因而丧失了其自我切割成熟的能力)相互作用的宿主蛋白,通过质谱分析出细胞蛋白Fascin-1。通过免疫共沉淀实验证实了△3CDpro及△3Cpro与Fascin在293T细胞内相互作用,但3Dpro却不能,进一步GSTpull-down实验也证实未缺失蛋白3CDpro和3CPro同样能在体外与Fascin蛋白直接相互作用。荧光共聚焦显微镜观察它们均定位于BHK-21细胞质中并且有很好的共定位。上调或下调Fascin蛋白表达能分别抑制或促进FMDV基因组复制,尤其是负链RNA的合成,而未对PV造成影响；FMDV感染早期(推测为复制起始时)能降低细胞内Fascin基因mRNA水平。
Foot-and-mouth disease (FMD) is one of the most highly contagious diseases of animals or humans, and the causative agent foot-and-mouth disease virus (FMDV) rapidly replicates and spreads within the infected animal, among in-contact susceptible animals, and by aerosol. FMDV is the prototype member of the Aphthovirus genus of the family Picornaviridae. The virus exists in the form of seven different serotypes:A, O, C, Asial, and South African Territories 1 (SAT1), SAT2, and SAT3, but a large number of subtypes have evolved within each serotype. Its genome is a single-stranded, positive-sense RNA which is over 8000 bases in length covalently bound at 5'-terminus to a 23-24 amino acid residue genome-linked protein, 3B. Replication of the FMDV genome RNA follows the same general pattern as replication of other members in the Picornaviridae family which occurs via a complementary negative strand RNA intermediate. To efficiently complete the life cycle of FMDV, not only viral proteins are involved, but also some host factors play important roles in most steps of viral infection, and identification of such host factors and their contributions has long been recognized as an important frontier.
1. Construction of FMDV full-length cDNA clone.
To stabilize FMDV genome and its genetical information, we successfully constructed type O FMDV full-length cDNA clone vector, pBluescript-FMDV. Basing on three incision enzyme sites within viral genome, four fragments were cloned into pBluescriptⅡKS(+) vector sequentially and then integrated which has artificial 25 poly(A) in 3'-terminal and 15 poly(C) within 5'UTR.
2. Establishment of a duplex quantitative real-time RT-PCR assay for simultaneous detection and quantitation of foot-and-mouth disease viral positive-stranded RNAs and negative-stranded RNAs.
Simplified, cost-effective, two-step duplex quantitative real-time RT-PCR assay was shown to detect and quantify foot-and-mouth disease virus positive-stranded RNAs and negative-stranded RNAs simultaneously for improved investigation of the state of virus infection and replication. The primers and Taqman probes were selected from the coding regions of 2B gene and 3D gene respectively, which have the least variations among serotypes. Cells infected acutely, tissue samples and single cell samples were used for evaluation of the assay. At the early stages of virus infection in vitro, the replication level reached a peak at 9 h.p.i. and the negative strands were detectable until 3 h.p.i. The kinetics of ratios of positive strands to negative strands (+RNA/-RNA) in vivo in the liver, kidney and spleen were similar, which demonstrated that the replication dynamics were similar in the three organs.55 single cell samples out of 187 were positive by both positive strands qPCR and negative strands qPCR, the ratios (+RNA/-RNA) ranged from 15.6 to 1463.4 which showed considerable difference among single cell samples, indicating that active viral replication differs greatly in single cells. A duplex quantitative real-time RT-PCR was validated as effective and reliable.
3. Interaction between FMDV 3CDpro and host factor Fascin-1 in BHK-21 cells.
Both virus-encoded proteins and cellular factors are involved in foot-and-mouth disease virus (FMDV) replication and pathogenesis. Tandem affinity purification was used in our study to purify cellular interactors of FMDV recombined 3 CD premature protein,△3CDpro (deletion of C-terminal 30 amino acids of 3C fragment in order to inactivate its auto-cleaving function), one of which was then identified as Fascin-1 by mass spectrum. The interaction between Fascin-1 and△3CDpro or△3Cpro rather than 3Dpro was conformed by co-immunoprecipitation assays. Results of confocal immunofluorescence analysis showed that they were well co-located in the cytoplasm of BHK-21 cells. GST pull down assay demonstrated further that the fascin directly interact with the 3CDpro as well as one of its mature protein 3Cpro in vitro. Overexpression or down-regulated expression of fascin-1 could inhibit or promote the replication of FMDV in BHK-21 rather than PV in Vero cells, especially synthesis of negative-strand RNAs. Furthermore, the levels of fascin mRNA decreased in early step of FMDV infection (maybe the initiation of replication).
引文
白兴文,李平花,孙普,李永亮,包慧芳,卢曾军,曹轶梅,郭建宏,刘在新不同长度poly(C)对口蹄疫病毒基因工程毒的毒力影响。微生物学报2008,48(12)：1654-1658。
李爽,张润祥,宋鸽,高明春,刘湘涛,王君伟牛Asia 1型口蹄疫病毒感染性克隆的构建。生物工程学报2009,25(11)：1621-1626。
张威,范振毅,陈光Fascin基因在恶性肿瘤中的表达及临床意义。吉林医学2010,31(4)：527-528。
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