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In SARS-CoV-2, this process is driven by interactions between viral structural proteins and the endoplasmic reticulum\u2013Golgi intermediate compartment (ERGIC) membrane. The membrane (M) protein, an embedded homodimer and the most abundant viral component, exists in two conformations: a compact \u201cshort\u201d form and an elongated \u201clong\u201d form. Although M is essential for virion assembly, how its conformations contribute to the generation and organization of the membrane curvature required for budding has remained unknown. Here, we used all-atom and Martini coarse-grained molecular dynamics simulations to show that individual M proteins can induce distinct membrane curvatures, depending on their conformation. The long form bends the membrane around its C-terminal, forming a valley-like depression, while the short form predominantly bends the membrane away from the C-terminal producing an anisotropic ridge. The induced curvatures correspond to the bulb and neck regions of a budding virion, respectively. Coarse-grained simulations of M protein pairs further reveal that curvature modulates long-range, membrane-mediated M\u2013M interactions, leading to repulsion between dissimilar conformations. Together, these results suggest that the long and short forms of M naturally segregate to shape the virion\u2019s bulb and neck, potentially facilitating genome encapsulation and membrane scission. This mechanism provides a physical basis for coronavirus budding and suggests that conformationally encoded curvature fields may represent a general principle underlying the formation of enveloped viruses.<\/jats:p>\n                <jats:sec>\n                  <jats:title>Significance Statement<\/jats:title>\n                  <jats:p>Enveloped viruses must bend host cell membranes to form new viral particles, but the driving force for membrane bending has been unclear. Using molecular dynamics simulations, we show that the SARS-CoV-2 membrane protein, the most abundant structural protein of the virus, drives membrane bending, and that its two natural conformations generate opposite signs of curvature which match the geometry of distinct regions of a budding virus. Moreover, these induced curvatures cause proteins in different conformations to repel one another providing a physical basis for their spatial segregation. Our results suggest that the membrane protein\u2019s conformations can contribute significantly to the membrane remodeling needed for virus assembly and release, revealing a simple mechanism that may apply to other enveloped viruses.<\/jats:p>\n                <\/jats:sec>","DOI":"10.64898\/2026.07.06.736641","type":"posted-content","created":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T05:15:21Z","timestamp":1783574121000},"source":"Crossref","is-referenced-by-count":1,"title":["SARS-CoV-2 membrane protein conformations induce distinct membrane curvatures"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8827-035X","authenticated-orcid":false,"given":"Joseph","family":"McTiernan","sequence":"first","affiliation":[{"name":"Department of Physics, University of California, Merced, Merced, CA 95343"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Roya","family":"Zandi","sequence":"additional","affiliation":[{"name":"Department of Physics and Astronomy, University of California, Riverside, Riverside, CA 92521"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael E.","family":"Colvin","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry, University of California, Merced, Merced, CA 95343"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9369-8780","authenticated-orcid":false,"given":"Ajay","family":"Gopinathan","sequence":"additional","affiliation":[{"name":"Department of Physics, University of California, Merced, Merced, CA 95343"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026071104303707000_2026.07.06.736641v1.1","doi-asserted-by":"publisher","DOI":"10.1038\/S41579-020-00468-6"},{"issue":"1","key":"2026071104303707000_2026.07.06.736641v1.2","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1146\/annurev-virology-110615-042301","article-title":"Structure, function, and evolution of coronavirus spike proteins","volume":"3","year":"2016","journal-title":"Annual Review of Virology"},{"key":"2026071104303707000_2026.07.06.736641v1.3","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-021-00418-x"},{"key":"2026071104303707000_2026.07.06.736641v1.4","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2006.11.027"},{"issue":"1","key":"2026071104303707000_2026.07.06.736641v1.5","doi-asserted-by":"crossref","first-page":"4629","DOI":"10.1038\/s41467-021-24887-y","article-title":"Correlative multi-scale cryo-imaging unveils sars-cov-2 assembly and egress","volume":"12","year":"2021","journal-title":"Nature Communications"},{"key":"2026071104303707000_2026.07.06.736641v1.6","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.74.11.4967-4978.2000"},{"key":"2026071104303707000_2026.07.06.736641v1.7","doi-asserted-by":"publisher","DOI":"10.1016\/j.jsb.2010.11.021"},{"key":"2026071104303707000_2026.07.06.736641v1.8","doi-asserted-by":"publisher","DOI":"10.1016\/bs.aivir.2016.08.005"},{"key":"2026071104303707000_2026.07.06.736641v1.9","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2019.08.031"},{"issue":"1","key":"2026071104303707000_2026.07.06.736641v1.10","doi-asserted-by":"crossref","first-page":"1936","DOI":"10.1038\/s41467-021-21953-3","article-title":"The sars-cov-2 nucleocapsid protein is dynamic, disordered, and phase separates with rna","volume":"12","year":"2021","journal-title":"Nature Communications"},{"issue":"1","key":"2026071104303707000_2026.07.06.736641v1.11","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1038\/s41467-020-20768-y","article-title":"The sars-cov-2 nucleocapsid phosphoprotein forms mutually exclusive condensates with rna and the membrane-associated m protein","volume":"12","year":"2021","journal-title":"Nature Communications"},{"key":"2026071104303707000_2026.07.06.736641v1.12","doi-asserted-by":"crossref","unstructured":"Bertrand Boson , Vincent Legros , Bingjie Zhou , Eglantine Siret , Cyrille Mathieu , Fran\u00e7ois-Lo\u00efc Cosset , Dimitri Lavillette , and Sol\u00e8ne Denolly . 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