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Two possible hydrogen bonding partnerships in the His37 tetrad of the M2 proton channel. (a) One of two pairs of His37 residues showing the low-barrier hydrogen bond model, in which the proton is transferred through the breaking and reforming of the intermonomer imidazole-imidazolium hydrogen bonds between His37 residues; (b) the water-His37 hydrogen bonding model, in which the His37 shuttles protons through imidazole ring reorientations and exchanging protons with water.
