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Using Pt GSTF8 from poplar as a model enzyme, we use biochemical, structural and computational methods to demonstrate that arGSTs employ a GSH-dependent Brnsted base mechanism for conversion of the LDOX product flavan-3,3,4-triol ( 4 ) or flavan-3-on-4-ol ( 7 ) into flav-2-en-3,4-diol ( 8-B4 ), an equilibrium form of cyanidin