Pathogenesis of Singapore grouper iridovirus (SGIV) and its immune control strategies
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摘要:
虹彩病毒(Iridovirus)是目前海水和淡水养殖鱼类最严重的病毒性病原之一，已从100多种鱼类中分离鉴定出该病毒。石斑鱼虹彩病毒(Singapore grouper iridovirus, SGIV)是从新加坡养殖的患病石斑鱼中分离鉴定的高致病性虹彩病毒，是虹彩病毒科Iridoviridae蛙病毒属Iridovirus 1种新的病毒。本文从以下几个方面对SGIV的研究进行综述：SGIV的形态、超微结构及其在石斑鱼细胞中的复制和装配过程；SGIV病毒基因组、转录组、囊膜蛋白质组及miRNAs的解析；SGIV感染宿主靶标组织的鉴定；病毒侵染的入侵方式、运动轨迹和胞内运输的实时追踪；SGIV感染宿主引起类凋亡的死亡机制及介导的信号通路的揭示；多种宿主免疫抗病基因对病毒感染的调节作用；多种SGIV的检测技术，包括基于抗体的流式细胞技术、微流控芯片检测技术、环介导等温扩增技术及核酸适配体检测方法等；SGIV的灭活疫苗、亚单位疫苗和DNA疫苗的研制等。以期为深入阐明SGIV感染致病机理奠定坚实的理论基础，为发展抗病毒对策提供技术支撑。
Abstract:Iridovirus is one of the most serious viral pathogens in marine and freshwater cultured fish. Up to now, iridoviruses have been isolated and identified from more than 100 fish species worldwide. Singapore grouper iridovirus (SGIV), a novel species of Ranavirus, was isolated from diseased grouper in Singapore. On the basis of establishing a virus-sensitive cell infection model, morphology, ultrastructure, replication and biochemical characterization of SGIV in grouper host cells were studied by electron microscopy and biochemical analysis. The molecular biological characterizations of SGIV, including viral genome, transcriptome, envelope proteome and viral microRNAs, were systematically analyzed by Omics analysis. The interactions between SGIV and host were investigated from many aspects, including identifying the target tissues of SGIV infection, tracking the single virus entry and transport, non-apoptosis cell death induced by SGIV infection, functions of host immune related genes in virus infection. Meanwhile, a variety of SGIV detection technologies have been developed, including antibody-based flow cytometry, microfluidic chip detection technology platform system, loop-mediated isothermal amplification (LAMP) and nucleic acid aptamer detection method. In addition, SGIV inactivated vaccine, subunit vaccine and DNA vaccine were developed. The results provide a theoretical basis for better understanding of the pathogenic mechanism of SGIV infection, and offer technical supports for the prevention and control of SGIV.
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图 1 SGIV病毒的分离、纯化及细胞内的复制增殖[3, 10-11]
a：患病石斑鱼肿大的脾脏；b：正常的石斑鱼脾细胞；c：SGIV感染引起脾细胞明显的细胞病变；d：感染细胞的超微结果观察，VM：病毒装配区，EC：空衣壳，MC：成熟核衣壳，NU：细胞核，标尺1 μm；e~g分别表示SGIV病毒粒子的出芽、释放及重感染邻近细胞，标尺为100 nm
Figure 1. Isolation and purification of SGIV and its replication in cells
a: The enlarged spleen of diseased grouper; b: Mock grouper spleen cells; c: The cytopathic effects induced by SGIV infection; d: Ultrastructural observation of infected cells, VM: viromatrix, EC: empty capsid, MC: matured nucleocapsid, NU: nuclesus, scale bar is 1 μm; e~g indicate budding, release and reinfection of SGIV particles respectively, scale bars are 100 nm
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图 2 SGIV病毒进入介导的内吞方式[32]
a~c：SGIV病毒进入依赖于网格蛋白和巨胞饮，而不依赖于小窝蛋白，标尺为10 μm；d：SGIV进入细胞的模式图
Figure 2. The endocytosis mediated by SGIV entry
a~c: SGIV enters grouper cells via the clathrin-mediated endocytic pathway and micropinocytosis, but not via caveola-dependent endocytosis, scale bars are 10 μm; d: Model entry route of SGIV into GS cells
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图 3 SGIV能显著激活MAPK信号通路中关键的信号分子[34, 36]
a：SGIV能激活ERK的磷酸化；b：SGIV感染对MAPK信号通路的激活情况及MAPK信号通路在病毒感染过程中的作用模式图，SGIV：新加坡石斑鱼虹彩病毒，MAPK：丝裂原活化蛋白激酶，MEK：MAPK激酶，ERK：细胞外调节蛋白激酶，JNK：c-Jun氨基末端激酶，MAPKAPK：MAPK激活蛋白激酶
Figure 3. SGIV infection significantly activates the key molecules in the MAPK signaling pathway
a: SGIV induces phosphorylation of ERK; b: Activation of MAPK signaling pathway induced by SGIV infection and the action of MAPK signaling pathway during virus infection, SGIV: Singapore grouper iridovirus, MAPK: mitogen-activated protein kinase, MEK: MAPK kinase, ERK: extracellular regulated protein kinases, JNK：c-Jun amino-terminal kinase, MAPKAPK: MAPK-activated protein kinase
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