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Previous studies, including our own, have identified SMCHD1 as a restriction factor that suppresses the replication of multiple viruses. Here, we reveal that the antiviral activity of SMCHD1 is dynamically regulated by two different post-translational modifications. SUMOylation of SMCHD1 promotes its association with the viral genome and enhances its antiviral activity. In contrast, during herpes simplex virus 1 (HSV-1) infection, the viral E3 ligase ICP0 induces SMCHD1 ubiquitination and proteasomal degradation, thereby relieving viral restriction. Loss of ICP0 stabilizes SMCHD1 and leads to marked accumulation of SUMOylated SMCHD1, rendering ICP0-deficient HSV-1 more sensitive to SMCHD1-mediated inhibition. Together, our findings uncover a reciprocal SUMO-ubiquitin regulatory mechanism that governs SMCHD1 antiviral activity and highlight a refined virus-host arms race centered on biphasic modification of a single restriction factor.<\/jats:p>","DOI":"10.1371\/journal.ppat.1014371","type":"journal-article","created":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T17:43:08Z","timestamp":1782322988000},"page":"e1014371","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["SUMOylation and ubiquitination reciprocally regulate SMCHD1 antiviral activity against herpes simplex virus 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Lilley","year":"2011","journal-title":"PLoS Pathog"},{"issue":"9","key":"ppat.1014371.ref038","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.00006-17","article-title":"Impaired STING Pathway in Human Osteosarcoma U2OS Cells Contributes to the Growth of ICP0-Null Mutant Herpes Simplex Virus","volume":"91","author":"T Deschamps","year":"2017","journal-title":"J Virol"},{"issue":"4","key":"ppat.1014371.ref039","doi-asserted-by":"crossref","first-page":"2052","DOI":"10.1128\/JVI.74.4.2052-2056.2000","article-title":"Herpes simplex virus ICP0 mutants are hypersensitive to interferon","volume":"74","author":"KL Mossman","year":"2000","journal-title":"J Virol"},{"issue":"29","key":"ppat.1014371.ref040","doi-asserted-by":"crossref","first-page":"21551","DOI":"10.1074\/jbc.M611041200","article-title":"Sumoylation of Oct4 enhances its stability, DNA binding, and transactivation","volume":"282","author":"F Wei","year":"2007","journal-title":"J Biol 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Gao","year":"2025","journal-title":"mBio"},{"issue":"8","key":"ppat.1014371.ref062","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1038\/s41556-023-01185-x","article-title":"STING is a cell-intrinsic metabolic checkpoint restricting aerobic glycolysis by targeting HK2","volume":"25","author":"L Zhang","year":"2023","journal-title":"Nat Cell Biol"},{"issue":"1","key":"ppat.1014371.ref063","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1128\/JVI.02635-14","article-title":"Identification of TRIM27 as a novel degradation target of herpes simplex virus 1 ICP0","volume":"89","author":"SE Conwell","year":"2015","journal-title":"J Virol"},{"issue":"44","key":"ppat.1014371.ref064","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.1211302109","article-title":"Nuclear IFI16 induction of IRF-3 signaling during herpesviral infection and degradation of IFI16 by the viral ICP0 protein","volume":"109","author":"MH Orzalli","year":"2012","journal-title":"Proc Natl Acad Sci U S 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