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OVERVIEW OF ANTIVIRAL DRUG DISCOVERY AND DEVELOPMENT
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design. By employing the CoV Mpro active site in an automatic ligand docking program to map a set of compounds selected from the NCI compound database on a precomputed grid of the protease surface, the nonpeptidyl natural product sabadinine 32 was identi ed as a hit.232 Note that selection of these lead compounds relied on homology models based on the structures of HCoV and TGEV. In cells, AG7088 and sabadinine failed to show activity against SARS-CoV233 and murine hepatitis virus (a mouse CoV),232 respectively. Molecular models can be improved based on the structural information on the SARS-CoV Mpro, which was published recently.234, 235 In studies with a cloned full-length Mpro and a truncated form containing only the catalytic domains, a cluster of serine residues near the active site cavity was shown to be susceptible to being targeted by compounds containing boronic acid (exempli ed by 33), providing attractive scaffolds for drug design.236
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A three-dimensional homologue model of the catalytic domain of SARS-CoV RdRp was built based on the polymerases of several RNA viruses.221 Analysis of the model suggested that active nucleoside analogues would have the C30 endo sugar puckering conformation and contain groups at the 20 and 30 positions that are capable of hydrogen bonding. SARS-CoV entry is mediated by the viral spike (S) protein interacting with a host receptor, angiotensin-converting enzyme 2 (ACE2), leading to membrane fusion and delivery of viral genome to the cytoplasm.237 239 Recent drug discovery research targeting the S protein includes (1) construction of a three-dimensional
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INHIBITORS OF FLAVIVIRUSES
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homologue model of the ACE2 structure;240 (2) identi cation of a peptide, CP-1, derived from the HR2 region in the S2 domain of SARS-CoV S protein to guide the design of fusion inhibitors;241 (3) identi cation of human monoclonal antibodies against the S1 domain of S protein;242 and (4) silencing S protein expression in cultured cells by RNA interference (RNAi).224,243 Other RNAi approaches targeting different sites on the genome were also reported.224,244,245
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JEV, DENV, WNV, Kyasanur Forest virus, and certain tickborne encephalitis viruses are in the Flavivirus genus of the Flaviviridae family. In general, the in vitro and in vivo activity of ribavirin against aviviruses is very weak119 (see Table 3.3). However, the combination of ribavirin and IFN-a in vitro resulted in signi cant synergistic interactions against YFV.89 In hamsters, ribavirin might be effective in the early treatment of YFV.88 On the other hand, it has been shown to worsen the disease caused by WNV.246 IFN-a showed promising results in small open trials,247 but IFN-a 2a did not affect the outcome in children with JE in a double-blind placebo-controlled trial.248 Flaviviral virions are composed of a lipid bilayer with two or more viral glycoproteins, whose fate and functions are governed by the process of N-linked glycosylation.249,250 Blockage of the trimming step of N-linked glycosylation by a glucosidase inhibitor, such as N-nonyl-deoxynojirimycin 34 (NN-DNJ), has been shown to suppress DENV-2 and JEV in a dose-dependent manner.251 In addition, in a lethal mouse model of JEV, oral dosing of NN-DNJ reduced the mortality rate.251
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The avivirus viral genome encodes a polyprotein that is processed into three structural (C, prM, and E) and seven nonstructural (NS1, NS2A, NS2B, NS3, NS4A, NS4B, and NS5) proteins.119,252,253 NS3 [with serine protease, 50 -RNA triphosphatase (50 -RTPase), nucleoside triphosphatase (NTPase), and RNA helicase activities] and NS5 [with methyltransferase (MTase) and RdRp activities] have been considered to be the optimal targets for antiviral agents.253,254 The NTPase/ helicase associated with NS3 protein of WNV has been puri ed255 and assayed, resulting in the identi cation of several ring systems imidazo[4,5-d]pyridazine (e.g., HMC-HO4256 35), imidazo[4,5-e][1,3]diazepine (e.g., 36257,258), benzimidazole (e.g., DRBT259 37), and benzotriazole (e.g., TBBT259 38) that could serve as starting templates for the development of new helicase-speci c antivirals.
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