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colon expression protein muc 2 crypt level epithelium goblet epithelial disease goblet cells coliti control treatment phase enterocyte mucin mtx treatment regenerative phase damage proliferation secretion intestine study surface epithelium synthesis surface dekker j dss treatment pp epithelium dss-induced number regenerative function segment paneth villu paneth cells epithelial proliferation einerhand awc dekker muc 2 levels factor marker einerhand tff 3 chapter dss-induced colitis section difference muc 2 secretion tissue effect appendix jejunum change onset day 4 antibody precursor alteration apoptosi tff 3 expression activity physiol buller duodenum epithelial cells model ileum sulfate muc 2 synthesis mucus layer np epithelium animal day 0 mucus dss-coliti 6.5 h enterocyte-specific