ACKR3 and malignant colon neoplasm: In other cell systems, p38 MAPK mediates MMP-9 upregulation in response to several stimuli, including TNF-α and SDF-1/CXCR7 in bladder and ovarian, respectively, carcinoma cells [48, 49], and clusterin/thrombospondin-1 in platelet-stimulated colon cancer [50].