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Mammalian orthoreoviruses (MRV) are non-fusogenic, nonenveloped, icosahedral, RNA viruses, containing a 10-segmented double-stranded RNA genome, belonging to the family Reoviridae. Infection with many mammalian orthoreovirus (MRV) strains results in shutoff of host, but not viral, protein synthesis via protein kinase R (PKR) activation and phosphorylation of translation initiation factor eIF2 . When cells are under stressful environments, such as heat shock, oxidative stress, nutritional starvation, and viral infection, several kinases (PKR, PERK, HRI, or GCN) are activated, which phosphorylate eIF2 ser), resulting in the formation of stress granules (SGs), discrete areas in the cytoplasm where cellular mRNAs are held in a translationally inactive state. We examined MRV-infected cells to characterize SG formation in response to MRV infection. We found SGs formed at early times following infection (2-6 h p.i.) in a manner dependent on phosphorylation of eIF2 . MRV induced SG formation in all four eIF2 kinase knockout cell lines, suggesting at least two kinases are involved in induction of SGs. Inhibitors of MRV disassembly prevented MRV-induced SG formation, indicating that viral uncoating is a required step for SG formation. Inactivation of MRV virions by ultraviolet (UV) light, or treatment of MRV-infected cells with the translational inhibitor, puromycin, did not prevent SG formation, suggesting that viral transcription and translation are not required for SG formation. Viral cores were found to colocalize with SGs, however, cores from UVinactivated virions did not associate with SGs, suggesting viral core particles are recruited into SGs in a process that requires the synthesis of viral mRNA. These results demonstrate that MRV particles induce SGs in a step following viral disassembly but preceding viral mRNA transcription, and that core particles are themselves recruited to SGs, suggesting the

Characterization of mammalian orthoreovirus (MRV) induced stress granules (SGs) and implications of eIF2α phosphorylation on viral translation