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However, the regulatory mechanisms that couple these processes are unknown. In zebrafish, notochord morphogenesis is a driver of axis extension through both posterior progenitor addition and anterior vacuolation. To elucidate how progenitor dynamics and vacuole-driven cell expansion interact to elongate the notochord, we generated a mathematical model linking progenitor addition rate to the expansion of cells from anterior to posterior to simulate vacuolation rate. Comparing this with empirical measurements, we find that progenitor incorporation, together with vacuolation, produces a linear gradient in nearest neighbour distance. We next explored the role of YAP\/TAZ in regulating progenitor addition in mutants for the YAP\/TAZ inhibitor vgll4b. We find that vgll4b expression and YAP activity are enriched in posterior midline progenitors. Loss of vgll4b elevates YAP signalling, enhances progenitor addition, restricts vacuole expansion, and \u2013 after a transient buffering phase \u2013 compromises anterior-posterior axis elongation. These results support a long-range feedback mechanism linking progenitor recruitment to vacuolation, enabling the notochord to balance cellular input with volumetric expansion, thereby maintaining tissue proportions.<\/jats:p>","DOI":"10.1242\/dev.205951","type":"journal-article","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T08:36:55Z","timestamp":1786696615000},"source":"Crossref","is-referenced-by-count":1,"title":["Coupling Vgll4b\/Yap-regulated posterior cell addition with anterior vacuolation enables robust notochord elongation"],"prefix":"10.1242","volume":"153","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9916-896X","authenticated-orcid":false,"given":"Carlos","family":"Camacho-Macorra","sequence":"first","affiliation":[{"name":"University of Cambridge 1 Department of Genetics , , Cambridge, CB2 3EH ,","place":["UK"]},{"name":"Selwyn College, University of Cambridge 2 , Cambridge, CB3 9DQ ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9866-4141","authenticated-orcid":false,"given":"Alberto S.","family":"Ceccarelli","sequence":"additional","affiliation":[{"name":"School of Biosciences, University of Nottingham 3 , Nottingham, LE12 5RD ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5841-8408","authenticated-orcid":false,"given":"Guillermo","family":"Serrano N\u00e1jera","sequence":"additional","affiliation":[{"name":"University of Cambridge 1 Department of Genetics , , Cambridge, CB2 3EH ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5169-5488","authenticated-orcid":false,"given":"Dillan","family":"Saunders","sequence":"additional","affiliation":[{"name":"University of Cambridge 1 Department of Genetics , , Cambridge, CB2 3EH ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-0162-2505","authenticated-orcid":false,"given":"Isabella","family":"Boesgaard","sequence":"additional","affiliation":[{"name":"University of Cambridge 1 Department of Genetics , , Cambridge, CB2 3EH ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0868-2507","authenticated-orcid":false,"given":"Osvaldo","family":"Chara","sequence":"additional","affiliation":[{"name":"School of Biosciences, University of Nottingham 3 , Nottingham, LE12 5RD ,","place":["UK"]},{"name":"Instituto de Tecnolog\u00eda, Universidad Argentina de la Empresa 4 , Buenos Aires C1073AAO ,","place":["Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7838-839X","authenticated-orcid":false,"given":"Benjamin","family":"Steventon","sequence":"additional","affiliation":[{"name":"University of Cambridge 1 Department of Genetics , , Cambridge, CB2 3EH ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,9,11]]},"reference":[{"key":"2026091110430695500_DEV205951C1","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1242\/dev.110.1.115","article-title":"The mechanics of notochord elongation, straightening and stiffening in the embryo of Xenopus laevis","volume":"110","author":"Adams","year":"1990","journal-title":"Development"},{"key":"2026091110430695500_DEV205951C2","doi-asserted-by":"publisher","first-page":"10005","DOI":"10.3390\/ijms251810005","article-title":"Wnt\/\u03b2-catenin signaling regulates Yap\/Taz activity during embryonic development in zebrafish","volume":"25","author":"Astone","year":"2024","journal-title":"Int. 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