[{"month":"04","article_type":"original","file":[{"file_id":"21901","date_created":"2026-05-21T06:05:49Z","success":1,"file_size":5603164,"content_type":"application/pdf","creator":"dernst","checksum":"a90e905968648ac4425c256de901e9c3","file_name":"2026_PhysicalReviewX_Drozdowski.pdf","relation":"main_file","date_updated":"2026-05-21T06:05:49Z","access_level":"open_access"}],"intvolume":"        16","_id":"21899","abstract":[{"lang":"eng","text":"Cell extrusion is an essential mechanism for controlling cell density in epithelial tissues. Another essential element of epithelia is curvature, which is required to achieve complex shapes, like in the lung or intestine. Here, we introduce a three-dimensional bubbly vertex model to study the interplay between extrusion and curvature. We find a generic cellular bulging instability at topological defects, which is much stronger than for standard vertex models. Analyzing cell shapes in three-dimensional imaging data of spherical mouse colon organoids, we infer that pentagonal cells have an increased basal interfacial tension, suggesting that cells at topological defects react to the different force conditions. Using the bubbly vertex model, we show that such basal tensions stabilize against the predicted instability and result in better cell shape control than tissue-scale mechanisms such as lumen pressure and spontaneous curvature. Our theory suggests that epithelial curvature naturally leads to bulged and extrusionlike cell shapes because the interfacial curvature of individual cells at the defects strongly amplifies buckling effected by tissue-scale topological defects in elastic sheets. Our results highlight the complex interplay of forces across scales in three-dimensional tissue organization."}],"publication":"Physical Review X","department":[{"_id":"EdHa"}],"DOAJ_listed":"1","volume":16,"date_updated":"2026-05-21T06:08:11Z","oa":1,"tmp":{"legal_code_url":"https://creativecommons.org/licenses/by/4.0/legalcode","short":"CC BY (4.0)","image":"/images/cc_by.png","name":"Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)"},"has_accepted_license":"1","doi":"10.1103/x82g-cq7n","publication_identifier":{"issn":["2160-3308"]},"user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","article_number":"021023","citation":{"short":"O.M. Drozdowski, B. Kocameşe-Tamgac𝚤, K.E. Boonekamp, M. Boutros, U.S. Schwarz, Physical Review X 16 (2026).","ama":"Drozdowski OM, Kocameşe-Tamgac𝚤 B, Boonekamp KE, Boutros M, Schwarz US. Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved epithelial sheets. <i>Physical Review X</i>. 2026;16(2). doi:<a href=\"https://doi.org/10.1103/x82g-cq7n\">10.1103/x82g-cq7n</a>","chicago":"Drozdowski, Oliver M, Büşra Kocameşe-Tamgac𝚤, Kim E. Boonekamp, Michael Boutros, and Ulrich S. Schwarz. “Cell Bulging and Extrusion in a Three-Dimensional Bubbly Vertex Model for Curved Epithelial Sheets.” <i>Physical Review X</i>. American Physical Society, 2026. <a href=\"https://doi.org/10.1103/x82g-cq7n\">https://doi.org/10.1103/x82g-cq7n</a>.","mla":"Drozdowski, Oliver M., et al. “Cell Bulging and Extrusion in a Three-Dimensional Bubbly Vertex Model for Curved Epithelial Sheets.” <i>Physical Review X</i>, vol. 16, no. 2, 021023, American Physical Society, 2026, doi:<a href=\"https://doi.org/10.1103/x82g-cq7n\">10.1103/x82g-cq7n</a>.","ista":"Drozdowski OM, Kocameşe-Tamgac𝚤 B, Boonekamp KE, Boutros M, Schwarz US. 2026. Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved epithelial sheets. Physical Review X. 16(2), 021023.","apa":"Drozdowski, O. M., Kocameşe-Tamgac𝚤, B., Boonekamp, K. E., Boutros, M., &#38; Schwarz, U. S. (2026). Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved epithelial sheets. <i>Physical Review X</i>. American Physical Society. <a href=\"https://doi.org/10.1103/x82g-cq7n\">https://doi.org/10.1103/x82g-cq7n</a>","ieee":"O. M. Drozdowski, B. Kocameşe-Tamgac𝚤, K. E. Boonekamp, M. Boutros, and U. S. Schwarz, “Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved epithelial sheets,” <i>Physical Review X</i>, vol. 16, no. 2. American Physical Society, 2026."},"file_date_updated":"2026-05-21T06:05:49Z","quality_controlled":"1","OA_type":"gold","ddc":["530"],"day":"30","type":"journal_article","scopus_import":"1","language":[{"iso":"eng"}],"date_published":"2026-04-30T00:00:00Z","issue":"2","publisher":"American Physical Society","publication_status":"published","acknowledgement":"O. M. D., M. B., and U.S. S. acknowledge support from the Max Planck School Matter to Life, with funding by the German Federal Ministry of Education and Research (BMBF), the Dieter Schwarz Foundation, and the Max Planck Society. M. B. and U.S. S. acknowledge support from the cluster of excellence 3DMM2O (EXC 2082/1-390761711 and EXC 2082/2-390761711) funded by the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation). The authors acknowledge the data storage service SDS@hd supported by the Ministry of Science, Research and the Arts Baden-Württemberg (MWK) and the DFG through Grant No. INST 35/1503-1 FUGG. For the publication fee we acknowledge financial support by Heidelberg University. O. M. D. thanks Edouard Hannezo for valuable discussions. U.S. S. is a member of the Interdisciplinary Center for Scientific Computing (IWR) at Heidelberg.","status":"public","date_created":"2026-05-20T14:35:57Z","OA_place":"publisher","author":[{"full_name":"Drozdowski, Oliver M","first_name":"Oliver M","last_name":"Drozdowski","id":"cd4ed792-b872-11ef-bb90-b7b3a3f62f75"},{"full_name":"Kocameşe-Tamgac𝚤, Büşra","first_name":"Büşra","last_name":"Kocameşe-Tamgac𝚤"},{"first_name":"Kim E.","full_name":"Boonekamp, Kim E.","last_name":"Boonekamp"},{"last_name":"Boutros","full_name":"Boutros, Michael","first_name":"Michael"},{"full_name":"Schwarz, Ulrich S.","first_name":"Ulrich S.","last_name":"Schwarz"}],"year":"2026","title":"Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved epithelial sheets","article_processing_charge":"Yes","oa_version":"Published Version"}]