Here, knockdown of integrin β1 or integrin α5 suppressed the phosphorylation levels of JNK and c-JUN obviously in HCC cells grown on higher stiffness substrate, also downregulated the expression of LOXL2 and c-JUN (Fig. 3b), demonstrating that higher matrix stiffness-activated JNK/c-JUN pathway was partially reversed. Here, LOXL2 is linked to hepatocellular carcinoma.