The intestinal epithelium is maintained by intestinal stem cells (ISCs), which produce post-mitotic absorptive and secretory epithelial cells. Initial fate specification toward enteroendocrine, goblet, and Paneth cell lineages requires the; transcription factor ATOH1, which regulates differentiation of the secretory cell lineage. However, less is known about the origin of tuft cells, which participate in type 2 immune; responses to parasite infections and appear to differentiate independently of ATOH1.