3.1.1.25 a 1,4-lactone + H2O = a 4-hydroxyacid - - 3.1.1.25 a 1,4-lactone + H2O = a 4-hydroxyacid upon substrate binding, the zinc-bridging hydroxide anion is effectively protected from re-protonation by the plug-like function of the acyl tails whereupon the ring-opening reaction can proceed. Subsequent protonation of the product's alkoxide group can then be accomplished by the intramolecular proton transfer from the intermediate state's carboxylic acid group where the resulting carboxylate group better accommodates the still-neighboring zinc ion 730750