This includes five clusters of cancer-associated fibroblasts (CAFs) consisting of a population of COL11A1+ CAFs (COL11A1, COL8A2) implicated in collagen metabolism 38, a population of LAMP5 CAFs defined by expression of LAMP5 and cystatins CST1 and CST2 implicated in promoting EMT 38, a population of PI16+ CAFs defined by expression of PI16 and PCOLCE2, a population of DPT+ CAFs defined by expression of DPT and CAPN6, as well as population of CA12+ CAFs defined by expression of CA12 and SLC2A1 which has been implicated in glycolysis metabolism and hypoxia 38. Here, CST1 is linked to cancer.