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Cartoon representation of inter-helical carboxylate-hydroxyl H-bonds and H-bond clusters. (A) The inter-helical carboxylate-hydroxyl H-bond may couple protonation change with changes in local protein dynamics (Bondar and Smith, 2017). (B) An internal carboxylate group and an external proton antenna cluster communicate via a water wire. (C) An interface H-bond cluster, as observed for spike protein S bound to ACE2 (Karathanou et al., 2020).
