{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,27]],"date-time":"2026-09-27T03:06:20Z","timestamp":1790478380230,"version":"4.1.0"},"reference-count":61,"publisher":"The Company of Biologists","issue":"6","license":[{"start":{"date-parts":[[2021,3,15]],"date-time":"2021-03-15T00:00:00Z","timestamp":1615766400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"crossref","award":["NRF-2018R1A2B3005113"],"award-info":[{"award-number":["NRF-2018R1A2B3005113"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2021,3,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>The actomyosin complex plays crucial roles in various life processes by balancing the forces generated by cellular components. In addition to its physical function, the actomyosin complex participates in mechanotransduction. However, the exact role of actomyosin contractility in force transmission and the related transcriptional changes during morphogenesis are not fully understood. Here, we report a mechanogenetic role of the actomyosin complex in branching morphogenesis using an organotypic culture system of mouse embryonic submandibular glands. We dissected the physical factors arranged by characteristic actin structures in developing epithelial buds and identified the spatial distribution of forces that is essential for buckling mechanism to promote the branching process. Moreover, the crucial genes required for the distribution of epithelial progenitor cells were regulated by YAP and TAZ through a mechanotransduction process in epithelial organs. These findings are important for our understanding of the physical processes involved in the development of epithelial organs and provide a theoretical background for developing new approaches for organ regeneration.<\/jats:p>","DOI":"10.1242\/dev.190785","type":"journal-article","created":{"date-parts":[[2021,3,3]],"date-time":"2021-03-03T18:55:52Z","timestamp":1614797752000},"source":"Crossref","is-referenced-by-count":17,"title":["A mechanogenetic role for the actomyosin complex in branching morphogenesis of epithelial organs"],"prefix":"10.1242","volume":"148","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4751-7125","authenticated-orcid":false,"given":"Jin Man","family":"Kim","sequence":"first","affiliation":[{"name":"Department of Oral Microbiology and Immunology, School of Dentistry and Dental Research Institute, Seoul National University, Seoul 08826, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2364-7160","authenticated-orcid":false,"given":"YoungJu","family":"Jo","sequence":"additional","affiliation":[{"name":"Department of Physics, Korea Advanced Institute of Science and Technology (KAIST), Daejeon 34141, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4292-9160","authenticated-orcid":false,"given":"Ju Won","family":"Jung","sequence":"additional","affiliation":[{"name":"Department of Dentistry, CHA Bundang Medical Center, CHA University School of Medicine, Seongnam 13496, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kyungpyo","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Physiology, School of Dentistry and Dental Research Institute, Seoul National University, Seoul 03080, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2021,3,23]]},"reference":[{"key":"2021042701195229600_DEV190785C1","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1016\/S1534-5807(02)00410-0","article-title":"Tube or not tube: remodeling epithelial tissues by branching morphogenesis","volume":"4","author":"Affolter","year":"2003","journal-title":"Dev. Cell"},{"key":"2021042701195229600_DEV190785C2","doi-asserted-by":"publisher","first-page":"1047","DOI":"10.1016\/j.cell.2013.07.042","article-title":"A mechanical checkpoint controls multicellular growth through YAP\/TAZ regulation by actin-processing factors","volume":"154","author":"Aragona","year":"2013","journal-title":"Cell"},{"key":"2021042701195229600_DEV190785C3","doi-asserted-by":"publisher","first-page":"pdb prot5558","DOI":"10.1101\/pdb.prot5558","article-title":"Organ culture and immunostaining of mouse embryonic kidneys","volume":"2011","author":"Barak","year":"2011","journal-title":"Cold Spring Harb. Protoc."},{"key":"2021042701195229600_DEV190785C4","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1186\/1471-2121-12-52","article-title":"Analysis of the role of Ser1\/Ser2\/Thr9 phosphorylation on myosin II assembly and function in live cells","volume":"12","author":"Beach","year":"2011","journal-title":"BMC Cell Biol."},{"key":"2021042701195229600_DEV190785C5","first-page":"674","article-title":"An iterative image registration technique with an application to stereo vision","author":"Bruce","year":"1981"},{"key":"2021042701195229600_DEV190785C6","doi-asserted-by":"publisher","first-page":"3286","DOI":"10.1101\/gad.11.24.3286","article-title":"Wnt signaling: a common theme in animal development","volume":"11","author":"Cadigan","year":"1997","journal-title":"Genes Dev."},{"key":"2021042701195229600_DEV190785C7","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1038\/ncb3191","article-title":"Integration of actin dynamics and cell adhesion by a three-dimensional, mechanosensitive molecular clutch","volume":"17","author":"Case","year":"2015","journal-title":"Nat. Cell Biol."},{"key":"2021042701195229600_DEV190785C8","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1016\/j.ydbio.2009.09.037","article-title":"Identification of a mechanochemical checkpoint and negative feedback loop regulating branching morphogenesis","volume":"336","author":"Daley","year":"2009","journal-title":"Dev. Biol."},{"key":"2021042701195229600_DEV190785C9","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1007\/978-1-59745-019-5_5","article-title":"Explant culture of mouse embryonic whole lung, isolated epithelium, or mesenchyme under chemically defined conditions as a system to evaluate the molecular mechanism of branching morphogenesis and cellular differentiation","volume":"633","author":"Del Moral","year":"2010","journal-title":"Methods Mol. Biol."},{"key":"2021042701195229600_DEV190785C10","doi-asserted-by":"publisher","first-page":"364","DOI":"10.1002\/jcp.1041430222","article-title":"Tissue plasminogen activator messenger RNA levels increase in cultured human endothelial cells exposed to laminar shear stress","volume":"143","author":"Diamond","year":"1990","journal-title":"J. Cell. Physiol."},{"key":"2021042701195229600_DEV190785C11","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1038\/nature10137","article-title":"Role of YAP\/TAZ in mechanotransduction","volume":"474","author":"Dupont","year":"2011","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C12","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1016\/bs.ctdb.2018.03.006","article-title":"Extracellular matrix (ECM) and the sculpting of embryonic tissues","volume":"130","author":"Dzamba","year":"2018","journal-title":"Curr. Top. Dev. Biol."},{"key":"2021042701195229600_DEV190785C13","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1038\/ncb3336","article-title":"Mechanical regulation of a molecular clutch defines force transmission and transduction in response to matrix rigidity","volume":"18","author":"Elosegui-Artola","year":"2016","journal-title":"Nat. Cell Biol."},{"key":"2021042701195229600_DEV190785C14","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1016\/j.cell.2006.06.044","article-title":"Matrix elasticity directs stem cell lineage specification","volume":"126","author":"Engler","year":"2006","journal-title":"Cell"},{"key":"2021042701195229600_DEV190785C15","doi-asserted-by":"publisher","first-page":"570","DOI":"10.1016\/j.devcel.2008.03.003","article-title":"Collective epithelial migration and cell rearrangements drive mammary branching morphogenesis","volume":"14","author":"Ewald","year":"2008","journal-title":"Dev. Cell"},{"key":"2021042701195229600_DEV190785C16","doi-asserted-by":"publisher","first-page":"1435","DOI":"10.1016\/j.celrep.2017.07.032","article-title":"The epithelial circumferential actin belt regulates YAP\/TAZ through nucleocytoplasmic shuttling of merlin","volume":"20","author":"Furukawa","year":"2017","journal-title":"Cell Rep."},{"key":"2021042701195229600_DEV190785C17","doi-asserted-by":"publisher","first-page":"1625","DOI":"10.1242\/bio.013672","article-title":"Epithelial topography for repetitive tooth formation","volume":"4","author":"Gaete","year":"2015","journal-title":"Biol. Open"},{"key":"2021042701195229600_DEV190785C18","doi-asserted-by":"publisher","first-page":"1038","DOI":"10.1038\/nature05014","article-title":"The emergence of geometric order in proliferating metazoan epithelia","volume":"442","author":"Gibson","year":"2006","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C19","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1016\/j.ceb.2011.06.002","article-title":"Dynamics of actomyosin contractile activity during epithelial morphogenesis","volume":"23","author":"Gorfinkiel","year":"2011","journal-title":"Curr. Opin. Cell Biol."},{"key":"2021042701195229600_DEV190785C20","doi-asserted-by":"publisher","first-page":"4452","DOI":"10.1242\/dev.096628","article-title":"Artificial three-dimensional niches deconstruct pancreas development in vitro","volume":"140","author":"Greggio","year":"2013","journal-title":"Development"},{"key":"2021042701195229600_DEV190785C21","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1126\/science.1235249","article-title":"Mechanics of epithelial tissue homeostasis and morphogenesis","volume":"340","author":"Guillot","year":"2013","journal-title":"Science"},{"key":"2021042701195229600_DEV190785C22","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1073\/pnas.1312076111","article-title":"Theory of epithelial sheet morphology in three dimensions","volume":"111","author":"Hannezo","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2021042701195229600_DEV190785C23","doi-asserted-by":"publisher","first-page":"1070","DOI":"10.1177\/0022034511405330","article-title":"Dynamics of salivary gland morphogenesis","volume":"90","author":"Harunaga","year":"2011","journal-title":"J. Dent. Res."},{"key":"2021042701195229600_DEV190785C24","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1359\/jbmr.2003.18.1.58","article-title":"A short pulse of mechanical force induces gene expression and growth in MC3T3-E1 osteoblasts via an ERK 1\/2 pathway","volume":"18","author":"Hatton","year":"2003","journal-title":"J. Bone Miner. Res."},{"key":"2021042701195229600_DEV190785C25","doi-asserted-by":"publisher","first-page":"1133","DOI":"10.1038\/ncb2124","article-title":"Tissue elongation requires oscillating contractions of a basal actomyosin network","volume":"12","author":"He","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"2021042701195229600_DEV190785C26","doi-asserted-by":"publisher","first-page":"56","DOI":"10.4161\/org.19813","article-title":"Extracellular matrix and cytoskeletal dynamics during branching morphogenesis","volume":"8","author":"Kim","year":"2012","journal-title":"Organogenesis"},{"key":"2021042701195229600_DEV190785C27","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1016\/j.devcel.2015.08.012","article-title":"Localized smooth muscle differentiation is essential for epithelial bifurcation during branching morphogenesis of the mammalian lung","volume":"34","author":"Kim","year":"2015","journal-title":"Dev. Cell"},{"key":"2021042701195229600_DEV190785C28","doi-asserted-by":"publisher","first-page":"7566","DOI":"10.1038\/s41598-018-25957-w","article-title":"Voltage-dependent Ca2+ channels promote branching morphogenesis of salivary glands by patterning differential growth","volume":"8","author":"Kim","year":"2018","journal-title":"Sci. Rep."},{"key":"2021042701195229600_DEV190785C29","doi-asserted-by":"publisher","first-page":"804","DOI":"10.1038\/nature07427","article-title":"Generation of a prostate from a single adult stem cell","volume":"456","author":"Leong","year":"2008","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C30","doi-asserted-by":"publisher","first-page":"860","DOI":"10.1038\/nrm2522","article-title":"Beyond polymer polarity: how the cytoskeleton builds a polarized cell","volume":"9","author":"Li","year":"2008","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2021042701195229600_DEV190785C31","doi-asserted-by":"publisher","first-page":"e21130","DOI":"10.7554\/eLife.21130","article-title":"YAP is essential for mechanical force production and epithelial cell proliferation during lung branching morphogenesis","volume":"6","author":"Lin","year":"2017","journal-title":"eLife"},{"key":"2021042701195229600_DEV190785C32","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1016\/j.stemcr.2013.10.013","article-title":"Combined KIT and FGFR2b signaling regulates epithelial progenitor expansion during organogenesis","volume":"1","author":"Lombaert","year":"2013","journal-title":"Stem Cell Rep."},{"key":"2021042701195229600_DEV190785C33","doi-asserted-by":"publisher","first-page":"1506","DOI":"10.1126\/science.1162783","article-title":"Patterning mechanisms of branched organs","volume":"322","author":"Lu","year":"2008","journal-title":"Science"},{"key":"2021042701195229600_DEV190785C34","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.neuron.2011.07.029","article-title":"Long-range neuronal circuits underlying the interaction between sensory and motor cortex","volume":"72","author":"Mao","year":"2011","journal-title":"Neuron"},{"key":"2021042701195229600_DEV190785C35","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1007\/s00424-007-0419-8","article-title":"Softening of the actin cytoskeleton by inhibition of myosin II","volume":"456","author":"Martens","year":"2008","journal-title":"Pfl\u00fcgers Arch \u2013 Eur. J. Physiol."},{"key":"2021042701195229600_DEV190785C36","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1016\/j.ceb.2010.07.010","article-title":"Conserved F-actin dynamics and force transmission at cell adhesions","volume":"22","author":"Maruthamuthu","year":"2010","journal-title":"Curr. Opin. Cell Biol."},{"key":"2021042701195229600_DEV190785C37","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1038\/s41586-018-0444-0","article-title":"RAP2 mediates mechanoresponses of the Hippo pathway","volume":"560","author":"Meng","year":"2018","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C38","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.ydbio.2004.07.022","article-title":"Spatiotemporal regulation of morphogenetic molecules during in vitro branching of the isolated ureteric bud: toward a model of branching through budding in the developing kidney","volume":"275","author":"Meyer","year":"2004","journal-title":"Dev. Biol."},{"key":"2021042701195229600_DEV190785C39","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1002\/dvdy.20237","article-title":"Control of basement membrane remodeling and epithelial branching morphogenesis in embryonic lung by Rho and cytoskeletal tension","volume":"232","author":"Moore","year":"2005","journal-title":"Dev. Dyn."},{"key":"2021042701195229600_DEV190785C40","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1016\/j.devcel.2014.06.012","article-title":"Parasympathetic innervation regulates tubulogenesis in the developing salivary gland","volume":"30","author":"Nedvetsky","year":"2014","journal-title":"Dev. Cell"},{"key":"2021042701195229600_DEV190785C41","doi-asserted-by":"publisher","first-page":"021005","DOI":"10.1115\/1.4032128","article-title":"On Buckling Morphogenesis","volume":"138","author":"Nelson","year":"2016","journal-title":"J. Biomech. Eng."},{"key":"2021042701195229600_DEV190785C42","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1038\/35001602","article-title":"Interaction between Wnt and TGF-\u03b2 signalling pathways during formation of Spemann's organizer","volume":"403","author":"Nishita","year":"2000","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C43","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1242\/dev.112.3.855","article-title":"Substitution for mesenchyme by basement-membrane-like substratum and epidermal growth factor in inducing branching morphogenesis of mouse salivary epithelium","volume":"112","author":"Nogawa","year":"1991","journal-title":"Development"},{"key":"2021042701195229600_DEV190785C44","doi-asserted-by":"publisher","first-page":"562","DOI":"10.1126\/science.1191880","article-title":"Btbd7 regulates epithelial cell dynamics and branching morphogenesis","volume":"329","author":"Onodera","year":"2010","journal-title":"Science"},{"key":"2021042701195229600_DEV190785C45","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1109\/TSMC.1979.4310076","article-title":"A threshold selection method from gray-level histograms","volume":"9","author":"Otsu","year":"1979","journal-title":"IEEE Trans. Syst. Man Cybern."},{"key":"2021042701195229600_DEV190785C46","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.ccr.2005.08.010","article-title":"Tensional homeostasis and the malignant phenotype","volume":"8","author":"Paszek","year":"2005","journal-title":"Cancer Cell"},{"key":"2021042701195229600_DEV190785C47","doi-asserted-by":"publisher","first-page":"1110","DOI":"10.1038\/nature09566","article-title":"Planar polarized actomyosin contractile flows control epithelial junction remodelling","volume":"468","author":"Rauzi","year":"2010","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C48","doi-asserted-by":"publisher","first-page":"e1003319","DOI":"10.1371\/journal.pcbi.1003319","article-title":"Cell-based multi-parametric model of cleft progression during submandibular salivary gland branching morphogenesis","volume":"9","author":"Ray","year":"2013","journal-title":"PLoS Comput. Biol."},{"key":"2021042701195229600_DEV190785C49","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1007\/978-1-59745-413-1_21","article-title":"ECM and FGF-dependent assay of embryonic SMG epithelial morphogenesis: investigating growth factor\/matrix regulation of gene expression during submandibular gland development","volume":"522","author":"Rebustini","year":"2009","journal-title":"Methods Mol. Biol."},{"key":"2021042701195229600_DEV190785C50","doi-asserted-by":"publisher","first-page":"12309","DOI":"10.1038\/ncomms12309","article-title":"A critical role for NF2 and the Hippo pathway in branching morphogenesis","volume":"7","author":"Reginensi","year":"2016","journal-title":"Nat. Commun."},{"key":"2021042701195229600_DEV190785C51","doi-asserted-by":"publisher","first-page":"4733","DOI":"10.1038\/s41598-018-23042-w","article-title":"Mechanical stretch induced transcriptomic profiles in cardiac myocytes","volume":"8","author":"Rys\u00e4","year":"2018","journal-title":"Sci. Rep."},{"key":"2021042701195229600_DEV190785C52","doi-asserted-by":"publisher","first-page":"876","DOI":"10.1038\/nature01712","article-title":"Fibronectin requirement in branching morphogenesis","volume":"423","author":"Sakai","year":"2003","journal-title":"Nature"},{"key":"2021042701195229600_DEV190785C53","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"2021042701195229600_DEV190785C54","doi-asserted-by":"publisher","first-page":"e23499","DOI":"10.7554\/eLife.23499","article-title":"The Hippo pathway effector YAP is an essential regulator of ductal progenitor patterning in the mouse submandibular gland","volume":"6","author":"Szymaniak","year":"2017","journal-title":"eLife"},{"key":"2021042701195229600_DEV190785C55","doi-asserted-by":"publisher","first-page":"342","DOI":"10.1126\/science.1204831","article-title":"Control of mitotic spindle angle by the RAS-regulated ERK1\/2 pathway determines lung tube shape","volume":"333","author":"Tang","year":"2011","journal-title":"Science"},{"key":"2021042701195229600_DEV190785C56","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1038\/s41556-018-0142-z","article-title":"YAP\/TAZ upstream signals and downstream responses","volume":"20","author":"Totaro","year":"2018","journal-title":"Nat. Cell Biol."},{"key":"2021042701195229600_DEV190785C57","doi-asserted-by":"publisher","first-page":"1229","DOI":"10.1101\/gad.888601","article-title":"TEAD\/TEF transcription factors utilize the activation domain of YAP65, a Src\/Yes-associated protein localized in the cytoplasm","volume":"15","author":"Vassilev","year":"2001","journal-title":"Genes Dev."},{"key":"2021042701195229600_DEV190785C58","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1083\/jcb.200806030","article-title":"Segregation and activation of myosin IIB creates a rear in migrating cells","volume":"183","author":"Vicente-Manzanares","year":"2008","journal-title":"J. Cell Biol."},{"key":"2021042701195229600_DEV190785C59","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1007\/s10911-013-9288-2","article-title":"Protocol: ex vivo culture of mouse embryonic mammary buds","volume":"18","author":"Voutilainen","year":"2013","journal-title":"J. Mammary Gland Biol. Neoplasia"},{"key":"2021042701195229600_DEV190785C60","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1177\/0022034517702342","article-title":"Piezo2 expression in mechanosensitive dental primary afferent neurons","volume":"96","author":"Won","year":"2017","journal-title":"J. Dent. Res."},{"key":"2021042701195229600_DEV190785C61","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1038\/ncb3292","article-title":"Locally excitable Cdc42 signals steer cells during chemotaxis","volume":"18","author":"Yang","year":"2016","journal-title":"Nat. Cell Biol."}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1242\/dev.190785","content-type":"unspecified","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.190785\/1818379\/dev190785.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.190785\/1818379\/dev190785.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,27]],"date-time":"2021-04-27T02:53:52Z","timestamp":1619492032000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/148\/6\/dev190785\/237852\/A-mechanogenetic-role-for-the-actomyosin-complex"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3,15]]},"references-count":61,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2021,3,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.190785","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"value":"0950-1991","type":"print"},{"value":"1477-9129","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,15]]},"article-number":"dev190785"}}