{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T01:36:19Z","timestamp":1790645779066,"version":"4.1.0"},"reference-count":44,"publisher":"The Company of Biologists","issue":"23","license":[{"start":{"date-parts":[[2024,12,1]],"date-time":"2024-12-01T00:00:00Z","timestamp":1733011200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"DOI":"10.13039\/501100007492","name":"Fondation Bettencourt Schueller","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100007492","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Learning Planet Institute"},{"name":"Emergence Sorbonne Universit\u00e9"},{"name":"Agence Nationale de la Recherche \u2018Young researchers\u2019 instrument","award":["BiSEV ANR-23-CE02-0018"],"award-info":[{"award-number":["BiSEV ANR-23-CE02-0018"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2024,12,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>While the impressive singing abilities of birds are made possible by the syrinx, the upper vocal system (i.e. trachea, larynx and beak) could also play a role in sound filtration. Yet, we still lack a clear understanding of the range of elongation this system can undertake, especially along the trachea. Here, we used biplanar cineradiography and X-ray reconstruction of moving morphology (XROMM) to record 15 species of cadaveric birds from 9 different orders while an operator moved the birds\u2019 heads in different directions. In all studied species, we found elongation of the trachea to be correlated with neck extension, and significantly greater (ranging from 18 to 48% for the whole motion; and from 1.4 to 15.7% for the singing positions) than previously reported on a live singing bird (3%). This elongation or compression was not always homogeneous along its entire length. Some specimens showed increased lengthening in the rostral part and others in both the rostral and caudal parts of the vocal tract. The diversity of elongation patterns shows that trachea elongation is more complex than previously thought. Since tracheal lengthening affects sound frequencies, our results contribute to our understanding of the mechanisms involved in complex communication signals, one of the amazing traits we share with birds.<\/jats:p>","DOI":"10.1242\/jeb.247945","type":"journal-article","created":{"date-parts":[[2024,10,18]],"date-time":"2024-10-18T06:56:42Z","timestamp":1729234602000},"source":"Crossref","is-referenced-by-count":6,"title":["The avian vocal system: 3D reconstruction reveals upper vocal tract elongation during head motion"],"prefix":"10.1242","volume":"227","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-8895-2249","authenticated-orcid":false,"given":"Morgane","family":"Fournier","sequence":"first","affiliation":[{"name":"INSERM, System Engineering and Evolution Dynamics, Universit\u00e9 Paris Cit\u00e9 1 , 75004 Paris ,","place":["France"]},{"name":"Learning Planet Institute 2 , 75004 Paris ,","place":["France"]},{"name":"Institute of Ecology and Evolution, Universit\u00e4t Bern 3 , 3012 Bern ,","place":["Switzerland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0803-9519","authenticated-orcid":false,"given":"Rachel","family":"Olson","sequence":"additional","affiliation":[{"name":"INSERM, System Engineering and Evolution Dynamics, Universit\u00e9 Paris Cit\u00e9 1 , 75004 Paris ,","place":["France"]},{"name":"Learning Planet Institute 2 , 75004 Paris ,","place":["France"]},{"name":"University of Akron 4 Department of Biology , , Akron, OH 44325 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5746-4621","authenticated-orcid":false,"given":"Sam","family":"Van Wassenbergh","sequence":"additional","affiliation":[{"name":"Universiteit Antwerpen 5 Department of Biologie , , 2610 Antwerpen ,","place":["Belgium"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9374-1291","authenticated-orcid":false,"given":"Pauline","family":"Provini","sequence":"additional","affiliation":[{"name":"INSERM, System Engineering and Evolution Dynamics, Universit\u00e9 Paris Cit\u00e9 1 , 75004 Paris ,","place":["France"]},{"name":"Learning Planet Institute 2 , 75004 Paris ,","place":["France"]},{"name":"UMR MECADEV 7179 CNRS\/Mus\u00e9um National d\u2018Histoire Naturelle, 75005 6 D\u00e9partement Adaptations du Vivant , Paris ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2024,12,6]]},"reference":[{"key":"2024120609100930900_JEB247945C1","doi-asserted-by":"publisher","first-page":"20200241","DOI":"10.1098\/rstb.2020.0241","article-title":"Phylogenetic signal in the vocalizations of vocal learning and vocal non-learning birds","volume":"376","author":"Arato","year":"2021","journal-title":"Phil. Trans. R. Soc. B"},{"key":"2024120609100930900_JEB247945C3","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1002\/jez.589","article-title":"X-ray reconstruction of moving morphology (XROMM): precision, accuracy and applications in comparative biomechanics research","volume":"313A","author":"Brainerd","year":"2010","journal-title":"J. Exp. Zool."},{"key":"2024120609100930900_JEB247945C4","doi-asserted-by":"publisher","first-page":"653","DOI":"10.1016\/j.anbehav.2010.12.015","article-title":"Singing direction as a tool to investigate the function of birdsong: an experiment on sedge warblers","volume":"81","author":"Brumm","year":"2011","journal-title":"Anim. Behav."},{"key":"2024120609100930900_JEB247945C5","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1111\/j.1474-919X.1899.tb01486.x","article-title":"On the trachea of the freckled duck of Australia (Stictonetta naevosa)","volume":"41","author":"Campbell","year":"1899","journal-title":"Ibis"},{"key":"2024120609100930900_JEB247945C6","first-page":"215","article-title":"Gross morphological and histological features of larynx, trachea and syrinx in Japanese quail. Anatomia, Histologia, Embryologia","volume":"36","author":"\u00c7evik-Demirkan","year":"2007","journal-title":"J. Veterinary Med. Ser. C"},{"key":"2024120609100930900_JEB247945C7","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/B978-012473070-0\/50005-0","article-title":"Vocal fighting and flirting: the functions of birdsong","volume-title":"Nature\u2018s Music: The Science of Birdsong","author":"Collins","year":"2004"},{"key":"2024120609100930900_JEB247945C8","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1002\/neu.10332","article-title":"Tracheal length changes during zebra finch song and their possible role in upper vocal tract filtering","volume":"59","author":"Daley","year":"2004","journal-title":"J. Neurobiol."},{"key":"2024120609100930900_JEB247945C9","first-page":"69","article-title":"Univariate metrics are not adequate to measure avian body size","volume":"107","author":"Freeman","year":"1990","journal-title":"The Auk"},{"key":"2024120609100930900_JEB247945C10","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1111\/j.1469-7998.1999.tb01020.x","article-title":"Acoustic exaggeration of size in birds by tracheal elongation: comparative and theoretical analyses","volume":"248","author":"Fitch","year":"1999","journal-title":"J. Zool."},{"key":"2024120609100930900_JEB247945C11","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1159\/000028474","article-title":"The phonetic potential of nonhuman vocal tracts: comparative cineradiographic observations of vocalizing animals","volume":"57","author":"Fitch","year":"2000","journal-title":"Phonetica"},{"key":"2024120609100930900_JEB247945C12","doi-asserted-by":"publisher","first-page":"258","DOI":"10.1016\/S1364-6613(00)01494-7","article-title":"The evolution of speech: a comparative review","volume":"4","author":"Fitch","year":"2000","journal-title":"Trends Cogn. Sci."},{"key":"2024120609100930900_JEB247945C13","doi-asserted-by":"publisher","first-page":"1005","DOI":"10.1121\/1.2159434","article-title":"Model for vocalization by a bird with distensible vocal cavity and open beak","volume":"119","author":"Fletcher","year":"2006","journal-title":"J. Acoust. Soc. Am."},{"key":"2024120609100930900_JEB247945C14","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1121\/1.424620","article-title":"Acoustics of avian vocal tract","volume":"105","author":"Fletcher","year":"1999","journal-title":"J. Acoust. Soc. Am."},{"key":"2024120609100930900_JEB247945C15","doi-asserted-by":"publisher","first-page":"6t1g1jx7q","DOI":"10.5061\/dryad.6t1g1jx7q","article-title":"The avian vocal system: 3D reconstruction reveals upper vocal tract elongation during head motion [Dataset]","author":"Fournier","year":"2024","journal-title":"Dryad"},{"key":"2024120609100930900_JEB247945C16","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1002\/jez.588","article-title":"Scientific rotoscoping: a morphology-based method of 3-D motion analysis and visualization","volume":"313A","author":"Gatesy","year":"2010","journal-title":"J. Exp. Zool."},{"key":"2024120609100930900_JEB247945C17","first-page":"230","article-title":"Integrative physiology of fundamental frequency control in birds","volume":"107","author":"Goller","year":"2013","journal-title":"J. Phys."},{"key":"2024120609100930900_JEB247945C18","doi-asserted-by":"publisher","first-page":"429","DOI":"10.2307\/2406936","article-title":"Tracheal dead space in the respiration of birds","volume":"25","author":"Hinds","year":"1971","journal-title":"Evolution"},{"key":"2024120609100930900_JEB247945C19","first-page":"58","volume-title":"Tracheal anatomy of the Anatidae and its taxonomic significance","author":"Johnsgard","year":"1961"},{"key":"2024120609100930900_JEB247945C20","doi-asserted-by":"publisher","first-page":"1256","DOI":"10.1111\/jeb.12397","article-title":"Migrate small, sound big: functional constraints on body size promote tracheal elongation in cranes","volume":"27","author":"Jones","year":"2014","journal-title":"J. Evol. Biol."},{"key":"2024120609100930900_JEB247945C21","doi-asserted-by":"publisher","first-page":"20220728","DOI":"10.1098\/rsif.2022.0728","article-title":"Realistic three-dimensional avian vocal model demonstrates how shape affect sound filtering (Passer domesticus)","volume":"20","author":"Kazemi","year":"2023","journal-title":"J. R. Soc. Interface"},{"key":"2024120609100930900_JEB247945C22","first-page":"105","volume":"4","author":"King","year":"1989","journal-title":"Larynx and Trachea. Form and Function in Birds"},{"key":"2024120609100930900_JEB247945C23","doi-asserted-by":"publisher","first-page":"e0163348","DOI":"10.1371\/journal.pone.0163348","article-title":"On the morphological description of tracheal and esophageal displacement and its phylogenetic distribution in Avialae","volume":"11","author":"Klingler","year":"2016","journal-title":"PLoS ONE"},{"key":"2024120609100930900_JEB247945C24","doi-asserted-by":"publisher","first-page":"3701","DOI":"10.1242\/jeb.145383","article-title":"Validation of XMALab software for marker-based XROMM","volume":"219","author":"Kn\u00f6rlein","year":"2016","journal-title":"J. Exp. 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Biol."},{"key":"2024120609100930900_JEB247945C29","first-page":"651","article-title":"The anatomy of the rings and muscles of the trachea of Gallus domesticus","volume":"99","author":"McLelland","year":"1965","journal-title":"J. Anat."},{"key":"2024120609100930900_JEB247945C31","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1242\/jeb.01378","article-title":"Vocal tract filtering and sound radiation in a songbird","volume":"208","author":"Nelson","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2024120609100930900_JEB247945C33","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1242\/jeb.064717","article-title":"Vocal tract articulation revisited: the case of the monk parakeet","volume":"215","author":"Ohms","year":"2012","journal-title":"J. Exp. Biol."},{"key":"2024120609100930900_JEB247945C34","doi-asserted-by":"crossref","first-page":"obab012","DOI":"10.1093\/iob\/obab012","article-title":"Regional tongue deformations during chewing and drinking in the pig","volume":"3","author":"Olson","year":"2021","journal-title":"Integr. Org. Biol."},{"key":"2024120609100930900_JEB247945C35","doi-asserted-by":"publisher","first-page":"8215","DOI":"10.1038\/s41598-020-64935-z","article-title":"XROMM and diceCT reveal a hydraulic mechanism of tongue base retraction in swallowing","volume":"10","author":"Orsbon","year":"2020","journal-title":"Sci. Rep."},{"key":"2024120609100930900_JEB247945C36","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1007\/s00359-008-0397-0","article-title":"Vocal tract motor patterns and resonance during constant frequency song: the white-throated sparrow","volume":"195","author":"Riede","year":"2009","journal-title":"J. Comp. Physiol. A. Neuroethol. Sens. Neural. Behav."},{"key":"2024120609100930900_JEB247945C37","doi-asserted-by":"publisher","first-page":"4025","DOI":"10.1242\/jeb.01256","article-title":"Inflation of the esophagus and vocal tract filtering in ring doves","volume":"307","author":"Riede","year":"2004","journal-title":"J. Exp. Biol."},{"key":"2024120609100930900_JEB247945C38","doi-asserted-by":"publisher","first-page":"5543","DOI":"10.1073\/pnas.0601262103","article-title":"Songbirds tune their vocal tract to the fundamental frequency of their song","volume":"103","author":"Riede","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2024120609100930900_JEB247945C39","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1007\/s00359-012-0768-4","article-title":"The acoustic effect of vocal tract adjustments in zebra finches","volume":"199","author":"Riede","year":"2013","journal-title":"J. Comp. Physiol. A Neuroethol. Sens. Neural Behav. Physiol."},{"key":"2024120609100930900_JEB247945C40","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pbio.2006507","article-title":"The evolution of the syrinx: An acoustic theory","volume":"17","author":"Riede","year":"2019","journal-title":"PLoS Biol."},{"key":"2024120609100930900_JEB247945C41","doi-asserted-by":"publisher","first-page":"105701","DOI":"10.1088\/1361-6501\/ab23e7","article-title":"Methods for characterization and optimisation of measuring performance of stereoscopic x-ray systems with image intensifiers","volume":"30","author":"Sanctorum","year":"2019","journal-title":"Measure. Sci. Technol."},{"key":"2024120609100930900_JEB247945C43","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1242\/jeb.182.1.147","article-title":"Kinematics of birdsong: Functional correlation of cranial movements and acoustic features in sparrows","volume":"182","author":"Westneat","year":"1993","journal-title":"J. Exp. Biol."},{"key":"2024120609100930900_JEB247945C44","first-page":"390","article-title":"Movements of the larynx during crowing in the domestic cock","volume":"103","author":"White","year":"1968","journal-title":"J. Anat."},{"key":"2024120609100930900_JEB247945C45","doi-asserted-by":"publisher","first-page":"3497","DOI":"10.1242\/jeb.204.20.3497","article-title":"Choreography of song, dance and bead movements in the zebra finch (Taeniopygia guttata)","volume":"204","author":"Williams","year":"2001","journal-title":"J. Exp. Biol."},{"key":"2024120609100930900_JEB247945C46","volume-title":"ggplot2, Elegant Graphics for Data Analysis","author":"Wickham","year":"2016","edition":"2nd edn"},{"key":"2024120609100930900_JEB247945C47","doi-asserted-by":"publisher","first-page":"419","DOI":"10.3390\/biology2010419","article-title":"Parsimony and model-based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals","volume":"2","author":"Yuri","year":"2013","journal-title":"Biology"},{"key":"2024120609100930900_JEB247945C48","doi-asserted-by":"publisher","first-page":"1085","DOI":"10.1242\/jeb.00208","article-title":"Leg morphology and locomotion in birds: requirements for force and speed during ankle flexion","volume":"206","author":"Zeffer","year":"2003","journal-title":"J. Exp. Biol."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.247945\/3571386\/jeb247945.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.247945\/3586335\/jeb247945.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.247945\/3586335\/jeb247945.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,12,6]],"date-time":"2024-12-06T04:10:49Z","timestamp":1733458249000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/227\/23\/jeb247945\/363347\/The-avian-vocal-system-3D-reconstruction-reveals"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,12,1]]},"references-count":44,"journal-issue":{"issue":"23","published-print":{"date-parts":[[2024,12,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.247945","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"value":"0022-0949","type":"print"},{"value":"1477-9145","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,12,1]]},"article-number":"jeb247945"}}