MMP9 and cancer: The MMP and lysine oxidase (LOX) family proteins are responsible for collagen remodeling in cancer.25,26 Overexpression of Sema3C markedly increased MMP2, MMP9, LOX, and LOXL2 levels, and treatment of MHCC-97L cells with rhSema3C elevated MMP2, LOX, and LOXL2 but not MMP9 (Supplementary Fig. 5a, b).