Domain architecture of the type II AAA+ proteins PEX1, PEX6, p97 and NSF. The two N-domains of PEX1 and PEX6 (N1 and N2) homologous to the N-domain of p97 and NSF (N-domain), and the two AAA+ domains (D1 and D2) are shown (left). Note that mammalian PEX1 is longer than the yeast protein, possessing a different spacing between its N-domains, it also has an extended C-terminal tail after the D2 domain (marked with asterisks). Atomic model of a p97 hexamer (right; PDB code 5C18, [138]) showing the common type II AAA+ structure consisting of one ring of D1 domains on top of another ring of D2 domains, and the six N-domains located on the side of the D1 ring. Note that this p97 structure was obtained in the presence of a non-hydrolysable ATP analogue and in the absence of substrate [138].