The abundant homohexameric AAA + ATPase p97 (also known as valosin-containing protein, VCP) is highly conserved from Dictyostelium discoideum to human and a pivotal factor of cellular protein homeostasis it catalyzes the unfolding proteins. Owing its fundamental function in quality control pathways, regulated by more than 30 cofactors, including UBXD family, whose members all carry an Ubiquitin...