Source: http://www.asmscience.org/content/book/10.1128/9781555816698.ch07
Timestamp: 2019-04-21 08:21:19+00:00

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This chapter reviews the viral and host players in the viral RNA replication process and the forms of picornavirus RNA utilized, as well as the RNA/protein complexes that facilitate viral RNA synthesis. Cleavage of poly(rC)-binding protein 2 (PCBP2) by 3CD contributes to a switch from translation to RNA replication for poliovirus, as full-length PCBP2 functions in viral translation but the truncated PCBP2 cleavage product can only function in RNA replication. The cis- acting replication element (CRE) is an RNA structure required for picornavirus RNA replication and was first discovered in the HRV14 genome by McKnight and Lemon. The picornavirus polymerase will homodimerize, oligomerize, and interact with the viral proteins 3AB, VPg, and 3CD. Although research efforts designed to elucidate the mechanisms of RNA replication utilized by picornaviruses have been comprehensive during the past several decades, there is still much to understand about the protein players and viral RNA sequences involved in replication. It is still unknown how 3Dpol can bind to sequences as disparate as the 3’ NCR/poly(A) tract of picornavirus genomic RNAs and those found at the 3’ ends of negative-strand RNA intermediates. Importantly, additional inhibitors of picornavirus replication that target specific players involved in RNA replication complex assembly, initiation, and chain elongation need to be developed as potential therapeutics against this important class of human and animal viruses.
Forms of viral RNA in a picornavirus-infected cell. Following entry and uncoating, the picornavirus genomic RNA is altered by the cleavage of VPg from its 5′ terminus by an unidentified host cellular enzyme termed unlinkase. Genomic picorna-virus RNA molecules lacking VPg are the templates for translation. These templates for translation also serve as the templates for negative-strand RNA synthesis, which results in a duplex of template and newly synthesized product RNA termed the RF. Negative-strand RNA molecules (perhaps derived from the RF) act as template for positive-strand RNA synthesis in RI complexes. The RI complexes have multiple positive-strand RNAs synthesized from a single negative-strand template, resulting in asymmetric levels of positive- versus negative-strand viral RNAs in the infected cell. The positive-strand viral RNA molecules can then serve as templates for additional rounds of translation or negative-strand RNA synthesis, or they are packaged into virions for subsequent infection of other host cells.
Simplified scheme of ribonucleoprotein complexes formed on genomic RNAs and that have been demonstrated to play a role in enterovirus RNA replication. In this example, viral (i.e., 3CD, 3AB, 3Dpol, and VPg) or host cell (PCBP and PABP) proteins are shown interacting with stem-loop I (also known as the cloverleaf), the CRE, or the 3′ poly(A) tract. The a, b, c, and d subelements for stem-loop I are shown, as are RNA secondary structures representing the CRE and the 3′ NCR of enterovirus genomic RNA. The IRES and protein-coding region are also indicated (not drawn to scale).
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 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V. 
 V.