芜菁邹缩病毒复制酶小亚基P28在重复侵染排斥中的功能研究
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摘要
重复侵染排斥(Superinfection exclusion)或同源排斥(Homologous interference)是指已经成功侵染的病毒能够排斥相同或者亲缘关系相近的病毒再次侵染的现象。重复侵染排斥现象在动物病毒和植物病毒中广泛存在,在农业生产中常用来防治某些植物病毒的危害,即预先接种了病毒温和株系的作物可以抵抗或者减轻与之亲源关系相近的强毒株系的侵染和危害,因此又被称为交义保护(Cross protection)。白1929年重复侵染排斥现象被发现以来,很多研究报道试图解释这一现象,但到目前为l止其分子机制尚不清楚。芜菁皱缩病毒(Turnip crinkle virus, TCV)是番茄丛矮病毒科(Tombusviridae)麝香石竹斑驳病毒属(Carmovirus)成员,属ss(+) RNA病毒,基因组大小为4054nt,可以系统侵染拟南芥、本生烟等植物。在本研究中,采用TCV和拟南芥侵染体系,尝试分析了TCV不同突变体病毒之间重复侵染排斥的分子机理。
为了探究TCV在侵染拟南芥过程中重复侵染排斥现象的分子机理,在TCV外壳蛋白编码框终止密码子之后分别整合9段不同的21nt核苷酸片段,构建了TCV病毒的9种突变体病毒。将这9种TCV突变体病毒重复侵染野生型和dcl2/dcl4突变株拟南芥,实验结果表明在TCV不同突变体侵染过程中存在着重复侵染排斥现象,而基因沉默机制并不是TCV突变体之间重复侵染排斥的主要原因。TCV和碎米荠褪绿斑点病毒(CCFV).香石竹斑驳病毒(CarMV)进行的重复侵染实验证明,TCV与亲缘关系近的CCFV病毒之间也存在着重复侵染排斥现象,但不排斥同源关系较远的CarMV。为了进一步明确TCV不同突变体病毒之间重复侵染排斥的关键蛋白,将瞬时表达TCV编码的各个蛋白的农杆菌与含TCV-GFP重组病毒的农杆菌混合后注射本生烟,结果证明TCV复制酶小亚基P28蛋白能够抑制TCV病毒的复制。进一步的实验证明,P28蛋白能够在植物细胞内形成大聚合物并排斥了TCV病毒的侵染,但瞬时表达的P28蛋白并不影响TCV病毒正常表达自己的复制酶小亚基。利用两种不同启动子分别瞬时表达P28GFP蛋白和P28Mcherry蛋白,证明P28具有捕获不同来源P28单体并形成聚合体的功能,这表明P28蛋白很可能是通过捕获TCV病毒表达的复制酶小亚基,使其不能发挥作用,从而抑制TCV病毒的复制。通过半变性SDS聚丙烯酰胺凝胶电泳实验,证明TCV复制酶小亚基P28在病毒侵染过程中也能够形成大的聚合物,这进一步佐证了P28蛋白是TCV重复侵染排斥现象的关键蛋白,在TCV侵染过程中存在两种不同的状态,其形成的不具有复制能力的聚合体能够捕捉二次侵染的TCV表达的P28,导致其无法进行复制。综上所述,TCV突变体之间存在重复侵染排斥,复制酶小亚基P28是这种重复侵染排斥的关键蛋白,通过形成聚合物捕捉二次侵染的TCV表达的P28排斥其侵染。
甜菜黑色焦枯病毒(Beet black scorch virus, BBSV)是番茄丛矮病毒科坏死病毒属(Necrovirus)成员,也是ss (+) RNA病毒,基因组大小为3644nt。本实验室前期的研究证明,BBSV外壳蛋白(Coat protein, CP)能够定位于细胞核,N末端4KRNKGGKKSR13碱性氨基酸富集区是其细胞核定位信号,对这些碱性氨基酸的突变会影响BBSV病毒粒体的形成以及其在本生烟中的系统运动。
为了深入研究外壳蛋白N末端碱性氨基酸在BBSV侵染过程中的功能,针对这些碱性氨基酸进行了一系列的点突变和缺失突变,并对这些突变体病毒的复制能力、病毒粒体稳定性和系统运动能力进行了检测。通过本生烟原生质体侵染实验,证明在外壳蛋白N末端引入的突变并不显著影响BBSV基因组RNA的复制和外壳蛋白的表达。通过纯化这些突变体的病毒粒体和进行RNase保护实验,证明这些突变体病毒粒体的形成和稳定性都不同程度的降低,说明外壳蛋白N末端的碱性氨基酸参与了BBSV病毒粒体装配和稳定性。外壳蛋白和基因组RNA体外结合实验显示,N末端的碱性氨基酸与病毒基因组RNA的结合有关,其中4KR5为关键氨基酸。通过侵染性实验,证明这些包装缺失突变体病毒系统侵染本生烟的能力也受到不同程度的影响。综上所述,BBSV外壳蛋白N末端的碱性氨基酸不仅是核定位信号,还是外壳蛋白和基因组RNA结合的关键区域,也影响BBSV病毒粒体的装配和稳定性,BBSV系统侵染本生烟需要完整的和稳定的病毒粒体。
Superinfection exclusion (SE) or Homologous interference is a phenomenon in which a preexisting viral infection prevents a secondary infection with the same or a closely related virus. SE has been described for various viruses, including important vertebrte, invertebrte, and plant pathogens. Indeed, SE, which is commonly referred to as "cross-protection", has also been used as a protective measure by purposely infecting plants with mild isolates of a virus to reduce infection and losses due to more severe isolates. Since its first report in1929, cross-protection has been explained in various ways, but unfortunately, its molecular mechanism is still unclear. In this report, I adopted the Turnip crinkle virus (TCV)-Arabidopsis system as a model to study cross-protection. TCV is a small icosahedral virus with a monopartite, positive sense RNA genome of4,054nucleotides (nt) that belongs to the Carmoviruss genus, in the Tombusviridae family and has an extensive host range including Arabidopsis and N. benthamiana.
To unravel the molecular mechanism of cross-protection, a series of TCV variants were designed by inserting21nt long oligo nucleotides of different sequences immediately after the CP open reading frame. By inoculating WT and dcl2/dcl4Arabidopsis plants with nine TCV variants, I found that one variant excluded other TCV derivatives during co-infection and showed that plant antiviral gene silencing does not play an important role in TCV cross-protection. Then super-inoculation experiments were conducted between TCV, CCFV(Cardamine chlorotic fleck virus) and CarMV (Carnation mottle virus), As expected from previous studies, I found that cross-protection only occurred closely related viruses. To determine whether exclusion was conferred by an individual TCV protein, I assessed the potential of each TCV protein to repress TCV replication by agrobacterium infiltrations. The results revealed that TCV-encoded P28, which is required for replication, exerts a highly specific and potent repression of TCV replication. By confocal microscopy observations, I was able to demonstrate that P28repressed TCV replication and formed multi-molecular aggregates as large as cell nuclei. Strikingly, the repression of TCV replication by P28did not abolish the translation of P28from the TCV genome. Then I adapted two different promoters for transcription of the mRNA of P28-GFP and P28-Mcherry fusion proteins. The results revealed that P28was able to interfere with P28translation from another source to inhibit TCV replication. Furthermore, semi-denatuirng gel electrophoresis results indicated that P28formed large, SDS-resistant aggregates during TCV infection. These results prompted the hypothesis in which P28plays opposite roles at different stages of the TCV multiplication cycle. First, P28functions in replication of the "protecting" virus early in infection and second, later in infection P28forms large nonfunctional aggregates that sequester P28encoded by subsequently infecting TCV to prevent replication of the second virus. In general, the first portion of my thesis succeeded in unraveling a mechanism for Cross-protection between TCV variants and demonstrated that P28plays an important role by forming large aggregates that inhibit subsequent invasion of different virus strains.
Beet black scorch virus (BBSV) is a small single-stranded, positive-sense RNA plant virus belonging to the genus Necrovirus, in the Tombusviridae family. The28nm icosahedral particle encapsidate the3,644nucleotide (nt) monopartite genomic RNA. In a previous study, we identified a motif4KRNKGGKKSR13rich in K and R residues, in the N terminus of the BBSV coat protein. This motif mediates nuclear localization of the CP, and our experiments lead to a model that the K/R-rich motif has an important role in BBSV long distance movement.
To further assess the functions of the basic amino acid residues in this motif, a series of mutants were generated by either replacing one or more basic amino acids with alanines (A) or deleting the complete motif. The resulting mutants were exhaustively examined for their replication competence, particle assembly, and systemic spread. First, replication of the mutants was tested in protoplasts, but none of the mutants compromised BBSV RNA replication appreciably. However, the mutants showed varying degrees of assembly defects and stability of BBSV virions, as determined by virus purification and RNase protection assays. These experiments demonstrated that all basic residues within the N-terminal region of the BBSV CP are required for ensure proper assembly of BBSV virions. Furthermore, I was able to determine that4KR5are the two most critical residues required for RNA-CP interactions and virion assembly. These results were further confirmed in vitro by assessing RNA-binding activities of the mutated CPs. I also demonstrated the indispensability of assembled virions for BBSV systemic infections in N. benthamiana by showing that defects in virion assembly/stability caused by the various CP mutants correlated with abundance of virus RNAs in systemically infected leaves. In summary, the N-terminal basic domain of BBSV CP is essential for nuclear localization, efficient CP-RNA interactions and virion stability, and intact virions are required for viral systemic spread.
为了探究TCV在侵染拟南芥过程中重复侵染排斥现象的分子机理,在TCV外壳蛋白编码框终止密码子之后分别整合9段不同的21nt核苷酸片段,构建了TCV病毒的9种突变体病毒。将这9种TCV突变体病毒重复侵染野生型和dcl2/dcl4突变株拟南芥,实验结果表明在TCV不同突变体侵染过程中存在着重复侵染排斥现象,而基因沉默机制并不是TCV突变体之间重复侵染排斥的主要原因。TCV和碎米荠褪绿斑点病毒(CCFV).香石竹斑驳病毒(CarMV)进行的重复侵染实验证明,TCV与亲缘关系近的CCFV病毒之间也存在着重复侵染排斥现象,但不排斥同源关系较远的CarMV。为了进一步明确TCV不同突变体病毒之间重复侵染排斥的关键蛋白,将瞬时表达TCV编码的各个蛋白的农杆菌与含TCV-GFP重组病毒的农杆菌混合后注射本生烟,结果证明TCV复制酶小亚基P28蛋白能够抑制TCV病毒的复制。进一步的实验证明,P28蛋白能够在植物细胞内形成大聚合物并排斥了TCV病毒的侵染,但瞬时表达的P28蛋白并不影响TCV病毒正常表达自己的复制酶小亚基。利用两种不同启动子分别瞬时表达P28GFP蛋白和P28Mcherry蛋白,证明P28具有捕获不同来源P28单体并形成聚合体的功能,这表明P28蛋白很可能是通过捕获TCV病毒表达的复制酶小亚基,使其不能发挥作用,从而抑制TCV病毒的复制。通过半变性SDS聚丙烯酰胺凝胶电泳实验,证明TCV复制酶小亚基P28在病毒侵染过程中也能够形成大的聚合物,这进一步佐证了P28蛋白是TCV重复侵染排斥现象的关键蛋白,在TCV侵染过程中存在两种不同的状态,其形成的不具有复制能力的聚合体能够捕捉二次侵染的TCV表达的P28,导致其无法进行复制。综上所述,TCV突变体之间存在重复侵染排斥,复制酶小亚基P28是这种重复侵染排斥的关键蛋白,通过形成聚合物捕捉二次侵染的TCV表达的P28排斥其侵染。
甜菜黑色焦枯病毒(Beet black scorch virus, BBSV)是番茄丛矮病毒科坏死病毒属(Necrovirus)成员,也是ss (+) RNA病毒,基因组大小为3644nt。本实验室前期的研究证明,BBSV外壳蛋白(Coat protein, CP)能够定位于细胞核,N末端4KRNKGGKKSR13碱性氨基酸富集区是其细胞核定位信号,对这些碱性氨基酸的突变会影响BBSV病毒粒体的形成以及其在本生烟中的系统运动。
为了深入研究外壳蛋白N末端碱性氨基酸在BBSV侵染过程中的功能,针对这些碱性氨基酸进行了一系列的点突变和缺失突变,并对这些突变体病毒的复制能力、病毒粒体稳定性和系统运动能力进行了检测。通过本生烟原生质体侵染实验,证明在外壳蛋白N末端引入的突变并不显著影响BBSV基因组RNA的复制和外壳蛋白的表达。通过纯化这些突变体的病毒粒体和进行RNase保护实验,证明这些突变体病毒粒体的形成和稳定性都不同程度的降低,说明外壳蛋白N末端的碱性氨基酸参与了BBSV病毒粒体装配和稳定性。外壳蛋白和基因组RNA体外结合实验显示,N末端的碱性氨基酸与病毒基因组RNA的结合有关,其中4KR5为关键氨基酸。通过侵染性实验,证明这些包装缺失突变体病毒系统侵染本生烟的能力也受到不同程度的影响。综上所述,BBSV外壳蛋白N末端的碱性氨基酸不仅是核定位信号,还是外壳蛋白和基因组RNA结合的关键区域,也影响BBSV病毒粒体的装配和稳定性,BBSV系统侵染本生烟需要完整的和稳定的病毒粒体。
Superinfection exclusion (SE) or Homologous interference is a phenomenon in which a preexisting viral infection prevents a secondary infection with the same or a closely related virus. SE has been described for various viruses, including important vertebrte, invertebrte, and plant pathogens. Indeed, SE, which is commonly referred to as "cross-protection", has also been used as a protective measure by purposely infecting plants with mild isolates of a virus to reduce infection and losses due to more severe isolates. Since its first report in1929, cross-protection has been explained in various ways, but unfortunately, its molecular mechanism is still unclear. In this report, I adopted the Turnip crinkle virus (TCV)-Arabidopsis system as a model to study cross-protection. TCV is a small icosahedral virus with a monopartite, positive sense RNA genome of4,054nucleotides (nt) that belongs to the Carmoviruss genus, in the Tombusviridae family and has an extensive host range including Arabidopsis and N. benthamiana.
To unravel the molecular mechanism of cross-protection, a series of TCV variants were designed by inserting21nt long oligo nucleotides of different sequences immediately after the CP open reading frame. By inoculating WT and dcl2/dcl4Arabidopsis plants with nine TCV variants, I found that one variant excluded other TCV derivatives during co-infection and showed that plant antiviral gene silencing does not play an important role in TCV cross-protection. Then super-inoculation experiments were conducted between TCV, CCFV(Cardamine chlorotic fleck virus) and CarMV (Carnation mottle virus), As expected from previous studies, I found that cross-protection only occurred closely related viruses. To determine whether exclusion was conferred by an individual TCV protein, I assessed the potential of each TCV protein to repress TCV replication by agrobacterium infiltrations. The results revealed that TCV-encoded P28, which is required for replication, exerts a highly specific and potent repression of TCV replication. By confocal microscopy observations, I was able to demonstrate that P28repressed TCV replication and formed multi-molecular aggregates as large as cell nuclei. Strikingly, the repression of TCV replication by P28did not abolish the translation of P28from the TCV genome. Then I adapted two different promoters for transcription of the mRNA of P28-GFP and P28-Mcherry fusion proteins. The results revealed that P28was able to interfere with P28translation from another source to inhibit TCV replication. Furthermore, semi-denatuirng gel electrophoresis results indicated that P28formed large, SDS-resistant aggregates during TCV infection. These results prompted the hypothesis in which P28plays opposite roles at different stages of the TCV multiplication cycle. First, P28functions in replication of the "protecting" virus early in infection and second, later in infection P28forms large nonfunctional aggregates that sequester P28encoded by subsequently infecting TCV to prevent replication of the second virus. In general, the first portion of my thesis succeeded in unraveling a mechanism for Cross-protection between TCV variants and demonstrated that P28plays an important role by forming large aggregates that inhibit subsequent invasion of different virus strains.
Beet black scorch virus (BBSV) is a small single-stranded, positive-sense RNA plant virus belonging to the genus Necrovirus, in the Tombusviridae family. The28nm icosahedral particle encapsidate the3,644nucleotide (nt) monopartite genomic RNA. In a previous study, we identified a motif4KRNKGGKKSR13rich in K and R residues, in the N terminus of the BBSV coat protein. This motif mediates nuclear localization of the CP, and our experiments lead to a model that the K/R-rich motif has an important role in BBSV long distance movement.
To further assess the functions of the basic amino acid residues in this motif, a series of mutants were generated by either replacing one or more basic amino acids with alanines (A) or deleting the complete motif. The resulting mutants were exhaustively examined for their replication competence, particle assembly, and systemic spread. First, replication of the mutants was tested in protoplasts, but none of the mutants compromised BBSV RNA replication appreciably. However, the mutants showed varying degrees of assembly defects and stability of BBSV virions, as determined by virus purification and RNase protection assays. These experiments demonstrated that all basic residues within the N-terminal region of the BBSV CP are required for ensure proper assembly of BBSV virions. Furthermore, I was able to determine that4KR5are the two most critical residues required for RNA-CP interactions and virion assembly. These results were further confirmed in vitro by assessing RNA-binding activities of the mutated CPs. I also demonstrated the indispensability of assembled virions for BBSV systemic infections in N. benthamiana by showing that defects in virion assembly/stability caused by the various CP mutants correlated with abundance of virus RNAs in systemically infected leaves. In summary, the N-terminal basic domain of BBSV CP is essential for nuclear localization, efficient CP-RNA interactions and virion stability, and intact virions are required for viral systemic spread.
引文
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