P4HA1 and neoplasm: The stiffness of tumor ECM is often associated with relatively high levels of crosslinked collagen (Type I), occurring due to the excessive activity of lysyl oxidase (LOX), as well as increased integrin signaling in the tumor microenvironment mediated by collagen modifying enzymes such as P4HA1, P4HA2, PLOD2, and LOX (Holback and Yeo, 2011).