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One corresponds roughly to the active site of Ec QueD (site 1), with the second site (site 2) more peripheral and solvent exposed, some ≈14 Å away and coordinated by the active site nucleophile, Cys23, an invariant Cys specific to QueD2s, Cys18 (this is a His in the other major cluster of QueD2 sequences), and two other poorly conserved residues derived from an adjacent subunit within a tetrameric ring