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A second aminoacid residue of mechanistic importance is Arg15, which is widely conserved among the known alpha class GSTs ( a value of the thiol group, and mutation of Arg15 to Ala or His decreases the steroid isomerase activity by two orders of magnitude, twice the effect of the Tyr9Phe mutation ( The mechanistic studies of human GST A1-1 were extended to human GST A3-3 following the discovery of the latter enzyme (Johansson and Mannervik, 2001)