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Left, bistable relationship between the degree of crypt opening and lumen inflation ratio: early lumen inflation (c ; (i)->(ii)) impairs long-term crypt budding morphogenesis, whereas lumen inflation after normal development (d ; (iii)->(iv)) has negligible effect (number of samples N = 4 in each scenario). Data are presented as mean values s.d. Right, representative snapshots of four types of crypt state: (i) day 3.5 bulged organoid; (ii) day 4 organoid that remained bulged due to long-term lumen inflation; (iii) day 4 budded organoid (unperturbed); and (iv) day 4 budded organoid after lumen inflation. Images are the maximum z projections of the organoids with DAPI and Lyz staining. Scale bars, 50 m.
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