Similarly, prolyl 4-hydroxylase subunit alpha 1 (P4HA1) and procollagen C-endopeptidase enhancer (PCOLCE) were identified as up-regulated in the cancer GEMMs (table S3), which both have known roles in collagen modification and folding (56) and therefore could be involved in promoting desmoplasia in PDAC development. The gene discussed is BMP1; the disease is cancer.