article published yes Oliver M Drozdowski author cd4ed792-b872-11ef-bb90-b7b3a3f62f75 Büşra Kocameşe-Tamgac𝚤 author Kim E. Boonekamp author Michael Boutros author Ulrich S. Schwarz author EdHa department 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. https://research-explorer.ista.ac.at/download/21899/21901/2026_PhysicalReviewX_Drozdowski.pdf application/pdfno American Physical Society2026 eng 2160-330810.1103/x82g-cq7n 162 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. 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. 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>. 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> 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>. 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> O.M. Drozdowski, B. Kocameşe-Tamgac𝚤, K.E. Boonekamp, M. Boutros, U.S. Schwarz, Physical Review X 16 (2026). 218992026-05-20T14:35:57Z2026-05-21T06:08:11Z