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We also show that water exchange in the S3 state is governed by a different equilibrium than in S2, and that the exchange of the fast substrate water in the S2 state is unaffected by Ca/Sr substitution. These findings support that (i) O5 is the slowly exchanging substrate water, with W2 being the only other option, and (ii) either W2 or W3 is the fast exchanging substrate. The three remaining possibilities for O-O bond formation in PSII are discussed.Interanionic H-bonds (IAHBs) are unfavourable interactions in the gas phase becoming favo
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