These included age-increasing pro-inflammatory genes in macrophages (Sox4)94, and signaling mediators in goblet cells (Fgfr2, Tcf7l2, Hif1a)95–97 and trophocytes (Grem1)98; many of which are involved in epithelial dedifferentiation95,96,99, inflammation97, and malignant transformation98,100,101, especially colitis-associated100,101 (Extended Data Table 11). Here, GREM1 is linked to colitis.