Representative MnII and FeII coordination geometries in class I RNRs. Class Ia Fe II/II2-β2 (PDB accession code 1PIY) and class Ib Fe II/II2-β2 (PDB accession code 3N38) structures were used to generate the FeII panels at top left. Class Ib Mn II/II2-β2 structures from the Ec (white, PDB accession code 3N37) and Bs (blue, PDB accession code 4DR0) enzymes were used to generate the MnII panels at top right. Note that the MnII sites in the class Ib β2 structures tend to exhibit higher coordination numbers with more strict adherence to octahedral geometry than the FeII coordination environments in the class Ia or Ib β2s. This trend is more apparent in the class Ib Ec Mn II/II2-β2 structure than in the class Ib Bs Mn II/II2-β2, and interestingly, the Ec protein binds MnII more tightly than the Bs homolog.9 Shown at bottom is a comparison to the coordination environments in class Ic β at site 1 (MnII, left) and site 2 (FeII, right). The binding sites in class Ic β2 resemble those in the MnII structures from class Ib β2s in that they are six-coordinate with exogenous water ligands, but the geometric distortion, particularly axial to the His ligand, is similar to Fe II/II2-β2 metal-binding interactions.