In hepatocellular carcinoma cells, matrix stiffness upregulates LOXL2 through the integrin β1/α5-JNK-AP1 signalling pathway [58], while in M2 macrophages matrix stiffness activates the integrin β5-FAK-MEK1/2-ERK1/2 pathway, resulting in HIF1 upregulation and, consequently, an increase in LOXL2 expression [59]. Here, LOXL2 is linked to hepatocellular carcinoma.