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Crystal structure of E. coli TSase-dUMP-H4folate complex (PDB ID 1KZI) showing the phosphate-binding site. All of the arginines presented are within hydrogen bond lengths from the phosphoester of dUMP (2.7–2.9 Å). Methylene C7 (turquoise) was built on C5 of dUMP to mimic the exocyclic methylene intermediate (compound D in Scheme 1). C6 of H4folate and C7 of methylene intermediate are the hydride donor and acceptor, respectively.
