《Table 1 The main host proteins that have been proven to interact with CMV-encoded proteins》

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《A Review on the Pathogenicity of Cucumber Mosaic Virus》

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Viruses can encode very limited number of proteins，so they rely on host-encoded proteins for replication.Table 1 shows the host proteins that have been proven to interact with CMV-encoded proteins.During the replication of CMV，1a protein is anchored first on the tonoplast or tonoplast proteins.Then 2a protein is also anchored on the tonoplast by interacting with 1a protein through the amino acid at position 126 at the N-terminal，to enable viral replication with the assistance of other proteins.After the replication is completed，2a protein is phosphorylated，and dissociates in the cytoplasm，participating in the infection of other viruses[5].Tcoi1protein of tobacco was screened via yeast two-hybrid system，and proven to be able to interact with the methyltransferase domain of1a protein.High level of Tcoi1 expression is beneficial to CMV infection，whereas reducing Tcoi1 expression inhibits CMV infection，indicating that Tcoi1 protein is able to methylate 1a protein，assisting the replication and transport of the virus[48].Tobacco Tcoi2 protein can also interact with and phosphorylate the methyltransferase domain of 1a protein，and it is assumed that the phosphorylation of 1a protein may affect the interaction between 1a protein and 2a protein or other host factors[49].In addition，there is an interaction between tobacco Bromodomain containing RNA binding protein（BRP1）and 1a protein.However，silencing BRP1 does not completely inhibit the replication of the virus，suggesting that the protein probably influences indirectly viral replication by affecting directly the stability of the virus replication complex[50].Formate dehydrogenase（FDH）and calreticulin-3 precursor（CRT3）which interact with 1a protein helicase were also isolated and identified from Capsicum annuum‘Bukang’by Choi et al.[51]，and they found that FDH-silenced plants showed local infection in inoculated leaves，but not in upper（systemic）leaves，and in CRT3-silenced plants，infection was not observed in either the inoculated or the upper leaves.Tobacco Tsip1 protein can interact with both 1a and 2a proteins.The level of viral RNA decreases in Tsip1-overexpressing transgenic tobacco plants，and continues to increase in Tsip1-silenced plants，suggesting that Tsip1 directly affects viral replication by complexing with 1a and 2a proteins[52].Tobacco calcineurin B-like interacting protein kinase（Nt CIPK12）that interacts with and phosphorylates 2a protein[53]，affecting the interaction of 2a with 1a[54].