{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T21:23:37Z","timestamp":1787779417486,"version":"build-2784847793"},"reference-count":122,"publisher":"Elsevier BV","issue":"8","license":[{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,7,22]],"date-time":"2026-07-22T00:00:00Z","timestamp":1784678400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001711","name":"Swiss National Science Foundation","doi-asserted-by":"publisher","award":["10007304"],"award-info":[{"award-number":["10007304"]}],"id":[{"id":"10.13039\/501100001711","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Cell Systems"],"published-print":{"date-parts":[[2026,8]]},"DOI":"10.1016\/j.cels.2026.101704","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T02:53:47Z","timestamp":1786589627000},"page":"101704","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["The emergence of the fractal bronchial tree"],"prefix":"10.1016","volume":"17","author":[{"given":"Malte","family":"Mederacke","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kevin A.","family":"Yamauchi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nikolaos","family":"Doumpas","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura","family":"Schaumann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan","family":"Sperl","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas","family":"Weikert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"D. Merrill","family":"Dane","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Connie C.W.","family":"Hsia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maurice","family":"Pradella","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jens","family":"Bremerich","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Roman","family":"Vetter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8051-1035","authenticated-orcid":false,"given":"Dagmar","family":"Iber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.cels.2026.101704_bib1","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1126\/science.137.3530.577","article-title":"Architecture of the human lung. Use of quantitative methods establishes fundamental relations between size and number of lung structures","volume":"137","author":"Weibel","year":"1962","journal-title":"Science"},{"key":"10.1016\/j.cels.2026.101704_bib2","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1152\/jappl.1968.24.3.373","article-title":"Morphology of the bronchial tree in man","volume":"24","author":"Horsfield","year":"1968","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib3","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1152\/japplphysiol.00838.2019","article-title":"The fractal geometry of bronchial trees differs by strain in mice","volume":"128","author":"Glenny","year":"2020","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib4","first-page":"1","article-title":"Das Prinzip des kleinsten Kraftverbrauchs im Dienste h\u00e4modynamischer Forschung","author":"Hess","year":"1914","journal-title":"Arch. Anat. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib5","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1073\/pnas.12.3.207","article-title":"The Physiological Principle of Minimum Work: I. The Vascular System and the Cost of Blood Volume","volume":"12","author":"Murray","year":"1926","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.cels.2026.101704_bib6","doi-asserted-by":"crossref","first-page":"668","DOI":"10.1038\/213668a0","article-title":"Design of the Bronchial Tree","volume":"213","author":"Wilson","year":"1967","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib7","series-title":"Lung Anatomy + Particle Deposition (lapd) Mouse Archive","author":"Beichel","year":"2019"},{"key":"10.1016\/j.cels.2026.101704_bib8","doi-asserted-by":"crossref","first-page":"1622","DOI":"10.1152\/japplphysiol.00562.2009","article-title":"Evidence for dysanapsis using computed tomographic imaging of the airways in older ex-smokers","volume":"107","author":"Sheel","year":"2009","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib9","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0132015","article-title":"Morphogenetic Implications of Peristalsis-Driven Fluid Flow in the Embryonic Lung","volume":"10","author":"Bokka","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.cels.2026.101704_bib10","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1016\/S1290-0729(00)01176-5","article-title":"Thermodynamic optimization of geometry: T- and Y-shaped constructs of fluid streams","volume":"39","author":"Bejan","year":"2000","journal-title":"Int. J. Therm. Sci."},{"key":"10.1016\/j.cels.2026.101704_bib11","series-title":"Morphometry of the Human Lung","author":"Weibel","year":"1963"},{"key":"10.1016\/j.cels.2026.101704_bib12","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/BF02476898","article-title":"Angles of branching and diameters of branches in the human bronchial tree","volume":"29","author":"Horsfield","year":"1967","journal-title":"Bull. Math. Biophys."},{"key":"10.1016\/j.cels.2026.101704_bib13","series-title":"Trachobronchial Geometry: Human, Dog, Rat, Hamster","author":"Raabe","year":"1976"},{"key":"10.1016\/j.cels.2026.101704_bib14","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1002\/ar.1091900202","article-title":"Application of an idealized model to morphometry of the mammalian tracheobronchial tree","volume":"190","author":"Phalen","year":"1978","journal-title":"Anat. Rec."},{"key":"10.1016\/j.cels.2026.101704_bib15","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1002\/ar.1091950308","article-title":"Anatomic models of the tracheobronchial and pulmonary regions of the rat","volume":"195","author":"Yeh","year":"1979","journal-title":"Anat. Rec."},{"key":"10.1016\/j.cels.2026.101704_bib16","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1016\/0034-5687(80)90050-X","article-title":"Branching angles in the bronchial tree related to order of branching","volume":"41","author":"Thurlbeck","year":"1980","journal-title":"Respir. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib17","doi-asserted-by":"crossref","first-page":"2310","DOI":"10.1152\/japplphysiol.00520.2004","article-title":"CT-based geometry analysis and finite element models of the human and ovine bronchial tree","volume":"97","author":"Tawhai","year":"2004","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib18","doi-asserted-by":"crossref","first-page":"L831","DOI":"10.1152\/ajplung.00281.2012","article-title":"Airway and pulmonary vascular measurements using contrast-enhanced micro-CT in rodents","volume":"304","author":"Counter","year":"2013","journal-title":"Am. J. Physiol., Lung Cell. Mol. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib19","doi-asserted-by":"crossref","DOI":"10.1103\/PhysRevE.93.062305","article-title":"Asymmetries arising from the space-filling nature of vascular networks","volume":"93","author":"Hunt","year":"2016","journal-title":"Phys. Rev. E"},{"key":"10.1016\/j.cels.2026.101704_bib20","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1152\/japplphysiol.00490.2017","article-title":"Morphological and functional properties of the conducting human airways investigated by in vivo computed tomography and in vitro MRI","volume":"124","author":"Van de Moortele","year":"2018","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib21","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1620\/tjem.194.157","article-title":"Irregularity of airway branching in a mouse bronchial tree: a 3-D morphometric study","volume":"194","author":"Onuma","year":"2001","journal-title":"Tohoku J. Exp. Med."},{"key":"10.1016\/j.cels.2026.101704_bib22","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1152\/jappl.1986.60.3.1089","article-title":"Beyond the principle of similitude: renormalization in the bronchial tree","volume":"60","author":"West","year":"1986","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib23","doi-asserted-by":"crossref","DOI":"10.1103\/PhysRevLett.106.178104","article-title":"Optimal branching asymmetry of hydrodynamic pulsatile trees","volume":"106","author":"Florens","year":"2011","journal-title":"Phys. Rev. Lett."},{"key":"10.1016\/j.cels.2026.101704_bib24","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1152\/japplphysiol.00984.2010","article-title":"An anatomical and functional model of the human tracheobronchial tree","volume":"110","author":"Florens","year":"2011","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib25","doi-asserted-by":"crossref","first-page":"168101","DOI":"10.1103\/PhysRevLett.95.168101","article-title":"Relating airway diameter distributions to regular branching asymmetry in the lung","volume":"95","author":"Majumdar","year":"2005","journal-title":"Phys. Rev. Lett."},{"key":"10.1016\/j.cels.2026.101704_bib26","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1016\/S0092-8240(77)80054-2","article-title":"Optimization of diameters and bifurcation angles in lung and vascular tree structures","volume":"39","author":"Uylings","year":"1977","journal-title":"Bull. Math. Biol."},{"key":"10.1016\/j.cels.2026.101704_bib27","article-title":"A generalized optimization principle for asymmetric branching in fluidic networks","volume":"472","author":"Stephenson","year":"2016","journal-title":"Proc. Math. Phys. Eng. Sci."},{"key":"10.1016\/j.cels.2026.101704_bib28","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/bs.ctdb.2021.02.002","article-title":"The control of lung branching morphogenesis","volume":"143","author":"Iber","year":"2021","journal-title":"Curr. Top. Dev. Biol."},{"key":"10.1016\/j.cels.2026.101704_bib29","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1038\/nature07005","article-title":"The branching programme of mouse lung development","volume":"453","author":"Metzger","year":"2008","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib30","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.devcel.2009.12.010","article-title":"Preparing for the first breath: genetic and cellular mechanisms in lung development","volume":"18","author":"Morrisey","year":"2010","journal-title":"Dev. Cell"},{"key":"10.1016\/j.cels.2026.101704_bib31","doi-asserted-by":"crossref","first-page":"L1082","DOI":"10.1152\/ajplung.00325.2016","article-title":"The total number of acini remains constant throughout postnatal rat lung development","volume":"311","author":"Barr\u00e9","year":"2016","journal-title":"Am. J. Physiol., Lung Cell. Mol. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib32","doi-asserted-by":"crossref","DOI":"10.1242\/dev.194209","article-title":"The biomechanical basis of biased epithelial tube elongation in lung and kidney development","volume":"148","author":"Conrad","year":"2021","journal-title":"Development"},{"key":"10.1016\/j.cels.2026.101704_bib33","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-018-05189-2","article-title":"Synchronized mesenchymal cell polarization and differentiation shape the formation of the murine trachea and esophagus","volume":"9","author":"Kishimoto","year":"2018","journal-title":"Nat. Commun."},{"key":"10.1016\/j.cels.2026.101704_bib34","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1161\/01.HYP.8.1.37","article-title":"Crucial role of endothelium in the vasodilator response to increased flow in vivo","volume":"8","author":"Pohl","year":"1986","journal-title":"Hypertension"},{"key":"10.1016\/j.cels.2026.101704_bib35","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1038\/nature01282","article-title":"Intracardiac fluid forces are an essential epigenetic factor for embryonic cardiogenesis","volume":"421","author":"Hove","year":"2003","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib36","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.78100","article-title":"Vein fate determined by flow-based but time-delayed integration of network architecture","volume":"12","author":"Marbach","year":"2023","journal-title":"eLife"},{"key":"10.1016\/j.cels.2026.101704_bib37","doi-asserted-by":"crossref","DOI":"10.1088\/1367-2630\/ad1488","article-title":"Vascular adaptation model from force balance: Physarum polycephalum as a case study","volume":"25","author":"Marbach","year":"2023","journal-title":"New J. Phys."},{"key":"10.1016\/j.cels.2026.101704_bib38","doi-asserted-by":"crossref","first-page":"1179","DOI":"10.1093\/aob\/mci130","article-title":"Biomechanical Model of the Xylem Vessels in Vascular Plants","volume":"95","author":"Karam","year":"2005","journal-title":"Ann. Bot."},{"key":"10.1016\/j.cels.2026.101704_bib39","first-page":"4328","article-title":"Microfluidic chest cavities reveal that transmural pressure controls the rate of lung development","volume":"144","author":"Nelson","year":"2017","journal-title":"Development"},{"key":"10.1016\/j.cels.2026.101704_bib40","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1152\/jappl.1996.81.1.209","article-title":"Regulation of lung expansion and lung growth before birth","volume":"81","author":"Harding","year":"1996","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib41","doi-asserted-by":"crossref","first-page":"77","DOI":"10.2976\/1.3043738","article-title":"Fluid dynamics in developmental biology: moving fluids that shape ontogeny","volume":"3","author":"Cartwright","year":"2009","journal-title":"HFSP J."},{"key":"10.1016\/j.cels.2026.101704_bib42","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1152\/jappl.1986.61.1.68","article-title":"Influence of upper respiratory tract on liquid flow to and from fetal lungs","volume":"61","author":"Harding","year":"1986","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib43","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1203\/00006450-199308000-00008","article-title":"Abolition of fetal breathing movements by spinal cord transection leads to reductions in fetal lung liquid volume, lung growth, and IGF-II gene expression","volume":"34","author":"Harding","year":"1993","journal-title":"Pediatr. Res."},{"key":"10.1016\/j.cels.2026.101704_bib44","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1085\/jgp.69.4.449","article-title":"Shear forces and blood vessel radii in the cardiovascular system","volume":"69","author":"Zamir","year":"1977","journal-title":"J. Gen. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib45","doi-asserted-by":"crossref","first-page":"35751","DOI":"10.1074\/jbc.M505832200","article-title":"Normal and cystic fibrosis airway surface liquid homeostasis. The effects of phasic shear stress and viral infections","volume":"280","author":"Tarran","year":"2005","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.cels.2026.101704_bib46","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1242\/dev.088500","article-title":"Spatial mapping and quantification of developmental branching morphogenesis","volume":"140","author":"Short","year":"2013","journal-title":"Development"},{"key":"10.1016\/j.cels.2026.101704_bib47","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0245558","article-title":"Bronchial tree of the human embryo: Categorization of the branching mode as monopodial and dipodial","volume":"16","author":"Fujii","year":"2021","journal-title":"PLoS One"},{"key":"10.1016\/j.cels.2026.101704_bib48","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1111\/joa.13946","article-title":"Bronchial tree of the human embryo: Examination based on a mammalian model","volume":"244","author":"Fujii","year":"2024","journal-title":"J. Anat."},{"key":"10.1016\/j.cels.2026.101704_bib49","doi-asserted-by":"crossref","first-page":"967","DOI":"10.1152\/jappl.1991.70.3.967","article-title":"Pleural pressure distribution and its relationship to lung volume and interstitial pressure","volume":"70","author":"Lai-Fook","year":"1991","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib50","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1152\/physrev.00026.2003","article-title":"Pleural mechanics and fluid exchange","volume":"84","author":"Lai-Fook","year":"2004","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.cels.2026.101704_bib51","unstructured":"Yartsev, A. (2019). Vertical gradient of pleural pressure. https:\/\/derangedphysiology.com\/main\/cicm-primary-exam\/required-reading\/respiratory-system\/Chapter%200356\/vertical-gradient-pleural-pressure."},{"key":"10.1016\/j.cels.2026.101704_bib52","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1113\/jphysiol.1981.sp013754","article-title":"Epithelial solute permeability, ion transport and tight junction morphology in the developing lung of the fetal lamb","volume":"315","author":"Olver","year":"1981","journal-title":"J. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib53","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1111\/j.1440-1681.1995.tb01988.x","article-title":"Fetal lung liquid: a major determinant of the growth and functional development of the fetal lung","volume":"22","author":"Hooper","year":"1995","journal-title":"Clin. Exp. Pharmacol. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib54","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1152\/jappl.1970.28.4.482","article-title":"Pressure drop and fluid flow regime of air inspired into the human lung","volume":"28","author":"Olson","year":"1970","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib55","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1002\/dvdy.70047","article-title":"Embryonic cerebrospinal fluid pressure in fetal mice in utero: External factors pressurize the intraventricular space","volume":"255","author":"Tsujikawa","year":"2025","journal-title":"Dev. Dyn."},{"key":"10.1016\/j.cels.2026.101704_bib56","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1152\/japplphysiol.00970.2001","article-title":"Dysanaptic growth of conducting airways after pneumonectomy assessed by CT scan","volume":"93","author":"Dane","year":"2002","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib57","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1152\/japplphysiol.01291.2013","article-title":"Defining a stimuli-response relationship in compensatory lung growth following major resection","volume":"116","author":"Ravikumar","year":"2014","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib58","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1152\/jappl.1957.10.1.1","article-title":"Influence of bronchial tree structure on ventilation in the dog\u2019s lung as inferred from measurements of a plastic cast","volume":"10","author":"Ross","year":"1957","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib59","doi-asserted-by":"crossref","first-page":"e1183","DOI":"10.1016\/S2214-109X(23)00217-6","article-title":"The global economic burden of chronic obstructive pulmonary disease for 204 countries and territories in 2020\u201350: a health-augmented macroeconomic modelling study","volume":"11","author":"Chen","year":"2023","journal-title":"Lancet Glob. Health"},{"key":"10.1016\/j.cels.2026.101704_bib60","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1164\/rccm.202301-0106PP","article-title":"Global Initiative for Chronic Obstructive Lung Disease 2023 Report: GOLD Executive Summary","volume":"207","author":"Agust\u00ed","year":"2023","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"10.1016\/j.cels.2026.101704_bib61","doi-asserted-by":"crossref","DOI":"10.1016\/j.ejrad.2022.110460","article-title":"Automated quantification of airway wall thickness on chest CT using retina U-Nets - Performance evaluation and application to a large cohort of chest CTs of COPD patients","volume":"155","author":"Weikert","year":"2022","journal-title":"Eur. J. Radiol."},{"key":"10.1016\/j.cels.2026.101704_bib62","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1111\/j.1469-7580.2007.00811.x","article-title":"Assessment of bronchial wall thickness and lumen diameter in human adults using multi-detector computed tomography: comparison with theoretical models","volume":"211","author":"Montaudon","year":"2007","journal-title":"J. Anat."},{"key":"10.1016\/j.cels.2026.101704_bib63","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1152\/jappl.1959.14.5.753","article-title":"Patterns of gas flow in the upper bronchial tree","volume":"14","author":"West","year":"1959","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib64","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1016\/0034-5687(70)90094-0","article-title":"The prediction of pressure drop and variation of resistance within the human bronchial airways","volume":"9","author":"Pedley","year":"1970","journal-title":"Respir. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib65","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.medengphy.2017.02.017","article-title":"N2 gas egress from patients\u2019 airways during LN2 spray cryotherapy","volume":"44","author":"O\u2019Connor","year":"2017","journal-title":"Med. Eng. Phys."},{"key":"10.1016\/j.cels.2026.101704_bib66","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1016\/0034-5687(85)90134-3","article-title":"Morphometric variability of the human upper bronchial tree","volume":"59","author":"Nikiforov","year":"1985","journal-title":"Respir. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib67","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1115\/1.3168381","article-title":"The Mechanism of Mucus Clearance in Cough","volume":"111","author":"Basser","year":"1989","journal-title":"J. Biomech. Eng."},{"key":"10.1016\/j.cels.2026.101704_bib68","series-title":"Mouse Embryogenesis: Methods and Protocols","first-page":"81","article-title":"Explant Culture for Studying Lung Development","volume":"1752","author":"Yeganeh","year":"2018"},{"key":"10.1016\/j.cels.2026.101704_bib69","doi-asserted-by":"crossref","first-page":"4526","DOI":"10.1242\/dev.116202","article-title":"An interplay of geometry and signaling enables robust lung branching morphogenesis","volume":"141","author":"Menshykau","year":"2014","journal-title":"Development"},{"key":"10.1016\/j.cels.2026.101704_bib70","doi-asserted-by":"crossref","first-page":"1439","DOI":"10.1126\/science.237.4821.1439","article-title":"Three-dimensional X-ray microtomography","volume":"237","author":"Flannery","year":"1987","journal-title":"Science"},{"key":"10.1016\/j.cels.2026.101704_bib71","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1016\/j.ejmp.2014.05.011","article-title":"Micro-CT of rodents: State-of-the-art and future perspectives","volume":"30","author":"Clark","year":"2014","journal-title":"Phys. Med."},{"key":"10.1016\/j.cels.2026.101704_bib72","series-title":"Der Bronchialbaum der S\u00e4ugethiere und des Menschen, nebst Bemerkungen \u00fcber den Bronchialbaum der V\u00f6gel und Reptilien","author":"Aeby","year":"1880"},{"key":"10.1016\/j.cels.2026.101704_bib73","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1004455","article-title":"Testing foundations of biological scaling theory using automated measurements of vascular networks","volume":"11","author":"Newberry","year":"2015","journal-title":"PLoS Comput. Biol."},{"key":"10.1016\/j.cels.2026.101704_bib74","unstructured":"Miettinen, A. (2025). pi2: C++ library and command-line software for processing and analysis of terabyte-scale volume images. v4.5. GitHub repository. https:\/\/github.com\/arttumiettinen\/pi2."},{"key":"10.1016\/j.cels.2026.101704_bib75","doi-asserted-by":"crossref","DOI":"10.3389\/fphys.2023.1302631","article-title":"Mechanosensitive channels in lung disease","volume":"14","author":"Zheng","year":"2023","journal-title":"Front. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib76","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s12013-009-9067-2","article-title":"Mechanotransduction by TRP channels: general concepts and specific role in the vasculature","volume":"56","author":"Yin","year":"2010","journal-title":"Cell Biochem. Biophys."},{"key":"10.1016\/j.cels.2026.101704_bib77","series-title":"PanVascular Medicine","first-page":"1","article-title":"Mechanotransduction and vascular resistance","author":"Thiriet","year":"2014"},{"key":"10.1016\/j.cels.2026.101704_bib78","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0040346","article-title":"Loss of the integrin-activating transmembrane protein Fam38A (Piezo1) promotes a switch to a reduced integrin-dependent mode of cell migration","volume":"7","author":"McHugh","year":"2012","journal-title":"PLoS One"},{"key":"10.1016\/j.cels.2026.101704_bib79","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1038\/nature21407","article-title":"Mechanical stretch triggers rapid epithelial cell division through Piezo1","volume":"543","author":"Gudipaty","year":"2017","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib80","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1186\/s12931-023-02328-2","article-title":"Intraluminal chloride regulates lung branching morphogenesis: involvement of piezo1\/piezo2","volume":"24","author":"Gon\u00e7alves","year":"2023","journal-title":"Respir. Res."},{"key":"10.1016\/j.cels.2026.101704_bib81","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1085\/jgp.78.4.431","article-title":"On connecting large vessels to small. The meaning of Murray\u2019s law","volume":"78","author":"Sherman","year":"1981","journal-title":"J. Gen. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib82","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1186\/s12915-021-01130-0","article-title":"Lepidoptera demonstrate the relevance of Murray\u2019s Law to circulatory systems with tidal flow","volume":"19","author":"Schachat","year":"2021","journal-title":"BMC Biol."},{"key":"10.1016\/j.cels.2026.101704_bib83","article-title":"Experimental models for Murray\u2019s law","volume":"50","author":"Akita","year":"2016","journal-title":"J. Phys. D Appl. Phys."},{"key":"10.1016\/j.cels.2026.101704_bib84","doi-asserted-by":"crossref","first-page":"939","DOI":"10.1038\/nature01444","article-title":"Water transport in plants obeys Murray\u2019s law","volume":"421","author":"McCulloh","year":"2003","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib85","doi-asserted-by":"crossref","first-page":"1017","DOI":"10.1161\/01.RES.77.5.1017","article-title":"Design principles of vascular beds","volume":"77","author":"Pries","year":"1995","journal-title":"Circ. Res."},{"key":"10.1016\/j.cels.2026.101704_bib86","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1007\/s11517-008-0330-2","article-title":"Wall shear stress as measured in vivo: consequences for the design of the arterial system","volume":"46","author":"Reneman","year":"2008","journal-title":"Med. Biol. Eng. Comput."},{"key":"10.1016\/j.cels.2026.101704_bib87","doi-asserted-by":"crossref","first-page":"1419","DOI":"10.1122\/1.4934240","article-title":"Murray\u2019s law revisited: Qu\u00e9mada\u2019s fluid model and fractal trees","volume":"59","author":"Moreau","year":"2015","journal-title":"J. Rheol."},{"key":"10.1016\/j.cels.2026.101704_bib88","doi-asserted-by":"crossref","first-page":"1476","DOI":"10.1161\/hy1201.100592","article-title":"Structural adaptation of vascular networks: role of the pressure response","volume":"38","author":"Pries","year":"2001","journal-title":"Hypertension"},{"key":"10.1016\/j.cels.2026.101704_bib89","first-page":"446","article-title":"Making microvascular networks work: angiogenesis, remodeling, and pruning","volume":"29","author":"Pries","year":"2014","journal-title":"Physiology (Bethesda)"},{"key":"10.1016\/j.cels.2026.101704_bib90","first-page":"3555","article-title":"Towards a unified theory for morphomechanics","volume":"367","author":"Taber","year":"2009","journal-title":"Philos. Trans. A Math. Phys. Eng. Sci."},{"key":"10.1016\/j.cels.2026.101704_bib91","doi-asserted-by":"crossref","first-page":"1646","DOI":"10.1016\/j.celrep.2015.02.035","article-title":"Axis Patterning by BMPs: Cnidarian Network Reveals Evolutionary Constraints","volume":"10","author":"Genikhovich","year":"2015","journal-title":"Cell Rep."},{"key":"10.1016\/j.cels.2026.101704_bib92","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1016\/j.cell.2017.08.026","article-title":"A Unifying Theory of Branching Morphogenesis","volume":"171","author":"Hannezo","year":"2017","journal-title":"Cell"},{"key":"10.1016\/j.cels.2026.101704_bib93","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1016\/j.cels.2024.08.001","article-title":"Versatile system cores as a conceptual basis for generality in cell and developmental biology","volume":"15","author":"Gallo","year":"2024","journal-title":"Cell Syst."},{"key":"10.1016\/j.cels.2026.101704_bib94","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.resp.2007.10.004","article-title":"Developmental regulation of lumenal lung fluid and electrolyte transport","volume":"159","author":"Wilson","year":"2007","journal-title":"Respir. Physiol. Neurobiol."},{"key":"10.1016\/j.cels.2026.101704_bib95","unstructured":"The Engineering ToolBox. (2014) Gases \u2013 dynamic (absolute) viscosities. https:\/\/www.engineeringtoolbox.com\/gases-absolute-dynamic-viscosity-d_1888.html."},{"key":"10.1016\/j.cels.2026.101704_bib96","series-title":"Proceedings of the Fourth Symposium on Underwater Physiology","first-page":"339","article-title":"Mechanics of breathing with helium\u2013oxygen and neon\u2013oxygen mixtures in deep saturation diving","author":"Anthonisen","year":"1971"},{"key":"10.1016\/j.cels.2026.101704_bib97","article-title":"Enabling peristalsis of human colon tumor organoids on microfluidic chips","volume":"14","author":"Fang","year":"2021","journal-title":"Biofabrication"},{"key":"10.1016\/j.cels.2026.101704_bib98","first-page":"1103","article-title":"Effect of tracheostomy on lung development in fetal lambs","volume":"55","author":"Fewell","year":"1983","journal-title":"J. Appl. Physiol. Respir. Environ. Exerc. Physiol."},{"key":"10.1016\/j.cels.2026.101704_bib99","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1038\/s41586-021-03416-3","article-title":"Ex utero mouse embryogenesis from pre-gastrulation to late organogenesis","volume":"593","author":"Aguilera-Castrejon","year":"2021","journal-title":"Nature"},{"key":"10.1016\/j.cels.2026.101704_bib100","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/S0925-4773(02)00451-3","article-title":"New insights into saccular development and vascular formation in lung allografts under the renal capsule","volume":"120","author":"Vu","year":"2003","journal-title":"Mech. Dev."},{"key":"10.1016\/j.cels.2026.101704_bib101","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1038\/s41592-023-02098-1","article-title":"The cousa objective: a long-working distance air objective for multiphoton imaging in vivo","volume":"21","author":"Yu","year":"2024","journal-title":"Nat. Methods"},{"key":"10.1016\/j.cels.2026.101704_bib102","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/1742-4682-6-7","article-title":"Extension of Murray\u2019s law using a non-Newtonian model of blood flow","volume":"6","author":"Revellin","year":"2009","journal-title":"Theor. Biol. Med. Model."},{"key":"10.1016\/j.cels.2026.101704_bib103","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1186\/1742-4682-3-31","article-title":"Pulsatile blood flow, shear force, energy dissipation and Murray\u2019s Law","volume":"3","author":"Painter","year":"2006","journal-title":"Theor. Biol. Med. Model."},{"key":"10.1016\/j.cels.2026.101704_bib104","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1016\/j.actbio.2022.10.047","article-title":"Biaxial mechanical properties of the bronchial tree: Characterization of elasticity, extensibility, and energetics, including the effect of strain rate and preconditioning","volume":"155","author":"Sattari","year":"2023","journal-title":"Acta Biomater."},{"key":"10.1016\/j.cels.2026.101704_bib105","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1152\/japplphysiol.01244.2010","article-title":"Lung tissue mechanics as an emergent phenomenon","volume":"110","author":"Suki","year":"2011","journal-title":"J. Appl. Physiol. (1985)"},{"key":"10.1016\/j.cels.2026.101704_bib106","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1016\/S0140-6736(71)91501-7","article-title":"Distribution of mechanical stress in the lung, a possible factor in localisation of pulmonary disease","volume":"1","author":"West","year":"1971","journal-title":"Lancet"},{"key":"10.1016\/j.cels.2026.101704_bib107","series-title":"Cardiovascular Mathematics: Modeling and Simulation of the Circulatory System","first-page":"177","article-title":"Chapter 5. Geometry and flow","author":"Doorly","year":"2009"},{"key":"10.1016\/j.cels.2026.101704_bib110","doi-asserted-by":"crossref","first-page":"1709","DOI":"10.1038\/nprot.2015.085","article-title":"Advanced CUBIC protocols for whole-brain and whole-body clearing and imaging","volume":"10","author":"Susaki","year":"2015","journal-title":"Nat. Protoc."},{"key":"10.1016\/j.cels.2026.101704_bib108","doi-asserted-by":"crossref","DOI":"10.1016\/j.xpro.2023.102187","article-title":"Time-lapse and cleared imaging of mouse embryonic lung explants to study three-dimensional cell morphology and topology dynamics","volume":"4","author":"G\u00f3mez","year":"2023","journal-title":"Star Protoc."},{"key":"10.1016\/j.cels.2026.101704_bib114","unstructured":"Haase, R., Yamauchi, K., M\u00fcller, J., and Fernando, I. (2022). haesleinhuepf\/apoc: 0.12.0. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.7489264."},{"key":"10.1016\/j.cels.2026.101704_bib122","unstructured":"Yamauchi, K., Haase, R., and Sobolewski, P. (2023). morphometrics\/morphometrics: v0.0.9. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.8256767."},{"key":"10.1016\/j.cels.2026.101704_bib112","doi-asserted-by":"crossref","first-page":"870","DOI":"10.1038\/s41592-019-0501-0","article-title":"BigStitcher: reconstructing high-resolution image datasets of cleared and expanded samples","volume":"16","author":"H\u00f6rl","year":"2019","journal-title":"Nat. Methods"},{"key":"10.1016\/j.cels.2026.101704_bib117","unstructured":"Schlegel, P., Gokaslan, A., and Kazimiers, T. (2024). skeletor. Software, version 1.3.0. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.10900419."},{"key":"10.1016\/j.cels.2026.101704_bib118","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.453","article-title":"scikit-image: image processing in python","volume":"2","author":"van der Walt","year":"2014","journal-title":"PeerJ"},{"key":"10.1016\/j.cels.2026.101704_bib120","unstructured":"Silversmith, W., Bae, J.A., Li, P.H., and Wilson, A.M. (2021). Kimimaro: Skeletonize densely labeled 3d image segmentations. Software, version 3.0.0. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.5539913."},{"key":"10.1016\/j.cels.2026.101704_bib121","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1056\/AIdbp2500994","article-title":"The TopCoW Challenge -- Topology-Aware Circle of Willis Segmentation for CT and MR Angiography","volume":"3","author":"Yang","year":"2026","journal-title":"NEJM AI"},{"key":"10.1016\/j.cels.2026.101704_bib111","doi-asserted-by":"crossref","first-page":"1105","DOI":"10.1038\/s41592-019-0554-0","article-title":"The mesoSPIM initiative: open-source light-sheet microscopes for imaging cleared tissue","volume":"16","author":"Voigt","year":"2019","journal-title":"Nat. Methods"},{"key":"10.1016\/j.cels.2026.101704_bib109","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1002\/dvg.20335","article-title":"A global double-fluorescent Cre reporter mouse","volume":"45","author":"Muzumdar","year":"2007","journal-title":"Genesis"},{"key":"10.1016\/j.cels.2026.101704_bib113","unstructured":"Ahlers, J., Althviz Mor\u00e9, D., Amsalem, O., Anderson, A., Bokota, G., Boone, P., Bragantini, J., Buckley, G., Burt, A., Bussonnier, M., et al. (2023). napari: a multi-dimensional image viewer for Python. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.8115575."},{"key":"10.1016\/j.cels.2026.101704_bib115","unstructured":"Comet Technologies Canada Inc. (2024). Dragonfly 2024.1. https:\/\/dragonfly.comet.tech."},{"key":"10.1016\/j.cels.2026.101704_bib116","doi-asserted-by":"crossref","unstructured":"Girshick, R. (2015). Fast r-cnn. Preprint at https:\/\/doi.org\/10.48550\/arXiv.1504.08083.","DOI":"10.1109\/ICCV.2015.169"},{"key":"10.1016\/j.cels.2026.101704_bib119","first-page":"462","article-title":"Building skeleton models via 3-D medial surface axis thinning algorithms","volume":"56","author":"Lee","year":"1994","journal-title":"CVGIP: Graph. Models Image Process."}],"container-title":["Cell Systems"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405471226001869?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405471226001869?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T21:13:35Z","timestamp":1787778815000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2405471226001869"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8]]},"references-count":122,"journal-issue":{"issue":"8","published-print":{"date-parts":[[2026,8]]}},"alternative-id":["S2405471226001869"],"URL":"https:\/\/doi.org\/10.1016\/j.cels.2026.101704","relation":{},"ISSN":["2405-4712"],"issn-type":[{"value":"2405-4712","type":"print"}],"subject":[],"published":{"date-parts":[[2026,8]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"The emergence of the fractal bronchial tree","name":"articletitle","label":"Article Title"},{"value":"Cell Systems","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.cels.2026.101704","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Author(s). Published by Elsevier Inc.","name":"copyright","label":"Copyright"}],"article-number":"101704"}}