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Transition state structure for Ras-catalyzed GTP hydrolysis as revealed by observed isotope effects. A charge δ (due to partial or full cleavage of the Oβ–γ–Pγ bond) is delocalized into the β phosphorus system. δ lies between –1 and –0.67 (0.02/0.03 = 0.67), which corresponds to an intrinsic KIE of 1.03, the maximum value observed for hydrolysis of phosphate monoesters, or of 1.02, the observed KIE, respectively.
