{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T10:00:15Z","timestamp":1792836015359,"version":"4.5.0"},"posted":{"date-parts":[[2026,9,18]]},"group-title":"Developmental Biology","reference-count":36,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T00:00:00Z","timestamp":1789689600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"award":["R56DE029124"],"award-info":[{"award-number":["R56DE029124"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"award":["R01DE018234"],"award-info":[{"award-number":["R01DE018234"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"award":["R01DE019638"],"award-info":[{"award-number":["R01DE019638"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"award":["F32DE018596"],"award-info":[{"award-number":["F32DE018596"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,9,18]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:sec>\n                  <jats:title>Background<\/jats:title>\n                  <jats:p>\n                    Successful palatal morphogenesis requires the precise spatiotemporal choreography of independent facial prominences. Our previous macroevolutionary analyses identified a convergence in amniote facial growth at primary palatal fusion, associated with reduced shape variability. We hypothesize that this constraint operates within individual species, representing a shared structural bottleneck where brain growth acts as a critical morphogenetic co-factor. To test this idea, we analyzed high-resolution 3D micro-CT images of embryonic chickens (\n                    <jats:italic>Gallus gallus<\/jats:italic>\n                    , N=217) spanning primary palatogenesis. Utilizing 3D geometric morphometrics (3DGM), we quantified a continuous \"developmental morphospace\" integrating the face and brain.\n                  <\/jats:p>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>Results<\/jats:title>\n                  <jats:p>Facial shape follows a highly nonlinear developmental trajectory. Using a continuous sliding-window analysis, we demonstrate that local morphological disparity undergoes a significant system-wide collapse coinciding with primary palatal fusion. Partial Least Squares (PLS) confirms this morphospace pivot is tightly coordinated with neurocranial development. Post-fusion, the modules uncouple: facial allometry switches from convergent to divergent projecting growth, while the brain undergoes spatial encapsulation, subsequently flatlining its morphological disparity.<\/jats:p>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>Conclusions<\/jats:title>\n                  <jats:p>Craniofacial morphogenesis can be described in terms of a tightly regulated biphasic developmental hourglass. The physical tethering of facial prominences creates a brief structurally constrained \"shape window\" essential for successful fusion. Deviations from this bottleneck \u2014 whether driven by altered prominence trajectories, increased local variance, or disruptions to the underlying forebrain landscape \u2014 push embryos into unoccupied morphospace, providing a unified mechanical and predictive framework for the spatiotemporal origins of non-syndromic cleft lip.<\/jats:p>\n                <\/jats:sec>","DOI":"10.64898\/2026.09.17.752413","type":"posted-content","created":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T21:10:24Z","timestamp":1789765824000},"source":"Crossref","is-referenced-by-count":1,"title":["Navigating the window: structural bottlenecks in avian craniofacial integration revealed by the ontogenetic patterning of shape variance"],"prefix":"10.64898","author":[{"given":"Diane","family":"Hu","sequence":"first","affiliation":[{"name":"Department of Orthopaedic Surgery, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jay","family":"Devine","sequence":"additional","affiliation":[{"name":"Department of Genetics and Electrical Engineering, KU Leuven, Leuven, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Cell Biology & Anatomy, University of Calgary, Calgary, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Benedikt","family":"Hallgr\u00edmsson","sequence":"additional","affiliation":[{"name":"Department of Cell Biology & Anatomy, University of Calgary, Calgary, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralph S.","family":"Marcucio","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3437-7493","authenticated-orcid":false,"given":"Nathan M.","family":"Young","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026092210301203000_2026.09.17.752413v1.1","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/2041-210X.12035","article-title":". geomorph: an R package for the collection and analysis of geometric morphometric shape data","volume":"4","year":"2013","journal-title":"Methods in Ecology and Evolution"},{"key":"2026092210301203000_2026.09.17.752413v1.2","doi-asserted-by":"publisher","DOI":"10.1666\/0094-8373-39.1.52"},{"key":"2026092210301203000_2026.09.17.752413v1.3","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2008.00999.x"},{"key":"2026092210301203000_2026.09.17.752413v1.4","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.20833"},{"key":"2026092210301203000_2026.09.17.752413v1.5","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.23726"},{"key":"2026092210301203000_2026.09.17.752413v1.6","first-page":"109","article-title":"Rapid three-dimensional phenotyping of cardiovascular development in mouse embryos by micro-CT with iodine staining","volume":"1","year":"2010","journal-title":"Circulation"},{"key":"2026092210301203000_2026.09.17.752413v1.7","doi-asserted-by":"publisher","DOI":"10.1007\/s11692-020-09508-8"},{"key":"2026092210301203000_2026.09.17.752413v1.8","doi-asserted-by":"crossref","unstructured":"Diewert VM , Lozanoff S . 2002. 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