Source: http://www.asmscience.org/content/book/10.1128/9781555817916.chap19
Timestamp: 2019-04-24 12:04:28+00:00

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Structure of poliovirus genomic RNA and processing of the polyprotein. The single-stranded RNA of poliovirus is shown with the terminal protein VPg at its 5′ end and the 3′ NTR with the poly(A) tail at its 3′ end. The 5′ NTR consists of the cloverleaf and the large IRES element. The location of the cre(2C) hairpin in the coding region of 2C is indicated. The attachment site of the 5′-terminal UMP to the tyrosine of VPg is shown enlarged. The polyprotein contains structural (P1) and nonstructural (P2, P3) domains. Processing of the P2 and P3 precursors of the polyprotein by 3Cpro/3CDpro is shown enlarged, with vertical lines indicating the proteinase cleavage sites.
Predicted secondary structures of picornaviral cis-replicating elements. (A) The PV1(M) 5′ cloverleaf. (B) The PV1(M) 3′ NTR-poly(A). (C) The PV1(M) cre(2C), HRV14 cre(VP1), and HRV2 cre(2A) RNAs. The conserved sequences in the loops are shown with bold letters. Also shown (boxed in) is the conserved sequence in all the known internal cis-replicating elements of picornaviral RNAs. See Note Added in Proof.
Proposed model of picornaviral minus-strand RNA synthesis. An RNP complex formed around the 5′ cloverleaf interacts with the PABP bound to the 3′ NTR-poly(A) resulting in a circularized genome ( 51a ). Proteinase 3CDpro cleaves membrane-bound 3AB to yield VPg and 3A. 3Dpol, 3CDpro, and VPg form a complex with the cre RNA hairpin. The polymerase synthesizes VPgpU and VPgpUpU using the A1A2ACA sequence in the loop as template, and the complex is transferred to the 3′ end of the poly(A) tail. The VPg-linked precursors then serve as primer for 3Dpol during the elongation step, a reaction possibly stimulated by membrane-bound 3AB.
Slide-back model of VPgpUpU synthesis by PV1(M) 3Dpol. Proteins 3Dpol, 3CDpro, and VPg form a complex with the PV1(M) cre(2C) RNA hairpin. Using A, in the A1A2CA sequence of the loop as template, the complementary nucleotide is selected and 3Dpol catalyzes the formation of a phosphodiester bond between UMP and the hydroxyl group of tyrosine in VPg. VPgpU then slides back and hydrogen bonds with A2 and the second UMP is added on the A1 template nucleotide.
Comparison of protein-primed RNA and DNA synthesis. (A) Initiation of picornaviral minus-strand RNA synthesis and phage Φ29 DNA synthesis. The slide-back mechanism is used by the picornaviral RNA polymerase and by phage Φ29 DNA polymerase ( 80 ) for the synthesis of the dinucleotidylylated protein precursors. Details of the mechanism are described in the text. (B) Picornaviral minus-strand RNA synthesis and HBV cDNA synthesis. Both viral polymerases use an internal RNA hairpin as the template for the protein-priming reaction and the nucleotidylylated proteins are translocated to the 3′ end of the plus strand where they are elongated into the complementary strands ( 53 , 99 , 122 ). (C) Picornaviral plus-strand RNA synthesis and phage Φ29 DNA synthesis ( 80 ). The end of the double-stranded template is first unwound by the binding of proteins to the plus and minus strands. The viral polymerases use the 3′ end of their RNA/DNA strand as template for the nucleotidylylation teaction. The precursors are elongated into complementary RNA/DNA strands. RT, reverse transcriptase; DP, DNA polymerase; TP, terminal protein.
Proposed model of picornaviral plus-strand RNA synthesis. The end of the RF is unwound by the binding of PCBP2/3CDpro and 3AB/3CDpro to the plus strand and of 2C to the minus strand of the 5′ cloverleaf. 3CDpro catalyzes the cleavage of membrane-bound 3AB to 3A and VPg, and 3CDpro undergoes autoprocessing. The polymerase synthesizes VPgpUpU using the 3′-terminal two As of the minus strand as template. The precursors are elongated into plus strands by the polymerase, possibly using the stimulatory activity of membrane-bound 3AB.
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