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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 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
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