Organization of the S. smaragdinae SPS-1 active site. The active sites of SPS-1 (green), SPM-1 (pink, PDB ID 4BP0), and NDM-1 (blue, PDB ID 4EXS) are shown as an overlay (a). The active site of SPS-1 (b) identifies a water molecule that coordinates to Zn1 in place of a histidine from the canonical HxHxD motif and a second water molecule which completes the hydrogen bond network (black dashed lines) with Gly96 and Asn192. The active sites of SPM-1 (c) and NDM-1 (d) feature the full complement of expected ligands. Throughout, key residues are shown as sticks and identified by residue name and number, zinc ions are shown as spheres (SPS-1: grey, SPM-1: navy blue, NDM-1: purple).