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Protonation state II. (a,b) For visual clarity, the carboxylate hydrogen bonding interactions are separated between a and b. Labels top and bottom refer to the position of Arg285 relative to His298. (c) The carbonyl forms hydrogen bonds with Thr301 and Ser62. (d) The tail carbonyl and the tail H–N2 group hydrogen bond to H2O-5 and H2O-6, respectively.
