Rev7 dimer is the organizing center of Polζ holoenzyme. a, Rev7 dimer presents a novel head-to-tail arrangement (left) in comparison to other HORMA family members (right, PDB ID: 2V64). The C-terminal seatbelt region is highlighted in cyan. Both Rev7 subunits are in the closed state with Rev7A binding RBM1 and Rev7B binding RBM2. b, Both Rev7 subunits are required for assembly of the Polζ holoenzyme. Structure and cryo-EM density details of the Rev7 dimer interface with Rev3, Pol31 and Pol32N. RBM1 and RBM2 share a consecutive proline-proline motif that interacts with similar set of residues (Y27, F141, L54, Y57, for example) from both monomers. Interactions at the Rev7A and RIR interface that further stabilize the Rev7A:Rev7B dimer are shown. In addition, interactions of Rev7B with Pol31PDE–Pol32N depicted here aid in restricting the movement of Pol31–Pol32N subcomplex relative to Rev3.