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Protonation state I. (a) Hydrogen bonds are depicted between His298, Arg285, and H2O-1 and the carboxylate group of benzylpenicillin. (b) The stabilizing hydrogen bond between O1 and Thr301 is depicted. (c) Shows water molecules that stabilize O4 (H2O-2), the sulfur (H2O-3), and the tail H–N2 (H2O-4).
