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While the cohesin ring serves as the central executor, cohesion is regulated by a remarkably intricate multi\u2010layered network extending far beyond the core complex. This review synthesizes current understanding of cohesion by integrating classical cohesin\u2010centric regulation with emerging evidence implicating diverse non\u2010cohesin factors, including DNA and RNA helicases, spliceosome components, microtubule\u2010associated proteins, and post\u2010translational modifications. We also examine the often\u2010overlooked contribution of DNA catenation to sister chromatid association and discuss how kinase\u2010phosphatase networks, chaperones, and non\u2010coding RNAs coordinate cohesion establishment, maintenance, and dissolution. Throughout this review, we emphasize that the cohesin complex remains the central mediator; the additional layers discussed here modulate, stabilize, or complement its function rather than replacing it. By integrating these diverse layers into a unified framework, we highlight unresolved questions and provide a roadmap for future investigations into the molecular mechanisms that ensure genomic stability.<\/jats:p>","DOI":"10.1002\/cbin.70220","type":"journal-article","created":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T12:44:57Z","timestamp":1790858697000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Sister Chromatid Cohesion: A Multilayered Regulatory Network Beyond the Cohesin Ring"],"prefix":"10.1002","volume":"50","author":[{"ORCID":"https:\/\/orcid.org\/0009-0003-5718-2212","authenticated-orcid":false,"given":"Jianing","family":"Fan","sequence":"first","affiliation":[{"name":"Bioscience and Biomedical Engineering Thrust Hong Kong University of Science and Technology (Guangzhou) Guangzhou Guangdong 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