1. Introduction p97, also known as valosin-containing protein (VCP), belongs to the AAA+ (ATPase associated with various cellular activities) ATPase family []. p97 is a homohexamer and each subunit contains an N-domain, two ATPase domains (D1 and D2), and a short unstructured C-terminal tail []. The N-domain is flexible and can therefore engage in interactions with varied adaptor proteins, leading to subcellular localization of p97 and the recognition of different substrates [].