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While several recent crystallographic and computational studies indicate that the O\n                    <jats:sub>x<\/jats:sub>\n                    ligand is absent in the S\n                    <jats:sub>3<\/jats:sub>\n                    state, we show that it forms upon entry into the S\n                    <jats:sub>4<\/jats:sub>\n                    state through migration of the Ca\u2010bound water ligand W\n                    <jats:sub>3<\/jats:sub>\n                    into the cavity between Mn1 and Ca\n                    <jats:sup>2<\/jats:sup>\n                    <jats:sup>+<\/jats:sup>\n                    . From this intermediate, O\n                    <jats:sub>x<\/jats:sub>\n                    \u2500O5 bond formation proceeds via the OHC mechanism rather than the conventional oxo\u2013oxyl pathway. This mechanism leads to formation of a hydroperoxide intermediate as the lowest\u2010energy post\u2010coupling species, in which Mn1 is reduced to the III oxidation state. Jahn\u2013Teller distortion considerably weakens the Mn1(III)\u2500O\n                    <jats:sub>x<\/jats:sub>\n                    bond, explaining the poor resolution of O\n                    <jats:sub>x<\/jats:sub>\n                    in XFEL structures following O\u2500O bond formation, an observation difficult to reconcile within conventional high\u2010oxidation\u2010state (HOS) models. Importantly, the proposed mechanism is consistent with water\u2010exchange experiments identifying W\n                    <jats:sub>3<\/jats:sub>\n                    and O5 as the substrate oxygen atoms. The mechanistic insights obtained in this work enabled us to propose likely S state structures and proton\u2010release assignments within the Kok cycle.\n                  <\/jats:p>","DOI":"10.1002\/chem.71765","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T03:47:33Z","timestamp":1791517653000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A Low\u2010Oxidation\u2010State Mechanism for O\u2500O Bond Formation in Photosystem II via Oxo\u2013Hydroxyl Radical Coupling"],"prefix":"10.1002","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2383-6380","authenticated-orcid":false,"given":"Alireza","family":"Ariafard","sequence":"first","affiliation":[{"name":"Research School of Chemistry Australian National University  Canberra 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