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Deamination of 5mC to thymine generates a G\/T mismatch, which occurs spontaneously and must be repaired prior to DNA replication to avoid mutation. We generated genetically engineered mice and cell lines to define DNA repair pathways that protect against 5mC deamination. We observed a low background mutation rate in mouse bone marrow or colon, typically 0.2\u20130.5 CG &amp;gt; TG mutations\/genome\/day. This increased 3\u20137 fold in cells lacking the glycosylase methyl-binding domain 4 (Mbd4), one of the few glycosylases capable of excising thymine from G\/T mismatches. We found no role for thymine DNA glycosylase (Tdg) in the repair of 5mC deamination in these tissues. Instead, our results flag potential cooperation between Mbd4 and the mismatch repair (MMR) complex MutS\u03b1 (Msh6:Msh2), evident through elevated rates of CG &amp;gt; TG mutations in Msh6-deficient cells: 2.6\u20134.8 CG &amp;gt; TG mutations\/genome\/day in primary cells and 13.9 CG &amp;gt; TG mutations\/genome\/day in cell lines. Loss of Mbd4 results in a specific increase in DNA damage from 5mC deamination, whereas the broader functionality of MutS\u03b1 explains why mutational signatures linked to Msh6 deficiency are more variable and reflect the replicative history of the cell. Our findings support the emerging view that MutS\u03b1 has broader DNA repair activity outside of replication.<\/jats:p>","DOI":"10.1093\/nar\/gkaf1493","type":"journal-article","created":{"date-parts":[[2025,12,19]],"date-time":"2025-12-19T12:52:50Z","timestamp":1766148770000},"source":"Crossref","is-referenced-by-count":3,"title":["Mbd4 and MutS\u03b1 protect cells from spontaneous deamination of 5-methylcytosine"],"prefix":"10.1093","volume":"54","author":[{"given":"Rebecca A","family":"Bilardi","sequence":"first","affiliation":[{"name":"Blood Cells and Blood Cancer Division, The Walter and Eliza Hall Institute , 1G Royal Pde, Parkville, VIC 3052 ,","place":["Australia"]},{"name":"Department of Medical Biology, University of Melbourne , 1G Royal Pde, Parkville, VIC 3052 ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christoffer","family":"Flensburg","sequence":"additional","affiliation":[{"name":"Blood Cells and Blood Cancer Division, The Walter and Eliza Hall Institute , 1G Royal Pde, Parkville, VIC 3052 ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhen","family":"Xu","sequence":"additional","affiliation":[{"name":"Blood Cells and Blood Cancer Division, The Walter and Eliza Hall Institute , 1G Royal Pde, Parkville, VIC 3052 ,","place":["Australia"]},{"name":"Department of Medical Biology, University of Melbourne , 1G Royal Pde, Parkville, VIC 3052 ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emily B","family":"Derrick","sequence":"additional","affiliation":[{"name":"Cancer Immunology Program, Peter MacCallum Cancer Centre , 305 Grattan St, Melbourne, VIC 3000 ,","place":["Australia"]},{"name":"Sir Peter MacCallum Department of Oncology, The University of Melbourne , Parkville, VIC 3010 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