Colonic epithelial cells were isolated in 10mM EDTA in DMEM, digested in a mixture of Accumax, Dispase, TrypLE , and sorted based on mCherry or GFP. RNA was extracted with Trizol reagent (ThermoFisher) for RNA-seq. Whole nuclei were used for chromatin transposition with Nextera Tn5 Transposase (Illumina) for ATAC-seq and ChiP-seq. SMARTer® Universal Low Input RNA Kit (TAKARA Bio) for RNA library and high-fidelity 2X PCR Master Mix (NEB) for ChIP and ATAC library were used.