{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T14:35:26Z","timestamp":1790951726421,"version":"4.1.0"},"reference-count":53,"publisher":"The Company of Biologists","issue":"14","license":[{"start":{"date-parts":[[2020,9,11]],"date-time":"2020-09-11T00:00:00Z","timestamp":1599782400000},"content-version":"vor","delay-in-days":2615,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2013,7,15]]},"abstract":"<jats:p>Syndecan 4 (Sdc4) is a cell-surface heparan sulfate proteoglycan (HSPG) that regulates gastrulation, neural tube closure and directed neural crest migration in Xenopus development. To determine whether Sdc4 participates in Wnt\/PCP signaling during mouse development, we evaluated a possible interaction between a null mutation of Sdc4 and the loop-tail allele of Vangl2. Sdc4 is expressed in multiple tissues, but particularly in the non-neural ectoderm, hindgut and otic vesicles. Sdc4;Vangl2Lp compound mutant mice have defective spinal neural tube closure, disrupted orientation of the stereocilia bundles in the cochlea and delayed wound healing, demonstrating a strong genetic interaction. In Xenopus, co-injection of suboptimal amounts of Sdc4 and Vangl2 morpholinos resulted in a significantly greater proportion of embryos with defective neural tube closure than each individual morpholino alone. To probe the mechanism of this interaction, we overexpressed or knocked down Vangl2 function in HEK293 cells. The Sdc4 and Vangl2 proteins colocalize, and Vangl2, particularly the Vangl2Lp mutant form, diminishes Sdc4 protein levels. Conversely, Vangl2 knockdown enhances Sdc4 protein levels. Overall HSPG steady-state levels were regulated by Vangl2, suggesting a molecular mechanism for the genetic interaction in which Vangl2Lp\/+ enhances the Sdc4-null phenotype. This could be mediated via heparan sulfate residues, as Vangl2Lp\/+ embryos fail to initiate neural tube closure and develop craniorachischisis (usually seen only in Vangl2Lp\/Lp) when cultured in the presence of chlorate, a sulfation inhibitor. These results demonstrate that Sdc4 can participate in the Wnt\/PCP pathway, unveiling its importance during neural tube closure in mammalian embryos.<\/jats:p>","DOI":"10.1242\/dev.091173","type":"journal-article","created":{"date-parts":[[2013,6,12]],"date-time":"2013-06-12T22:43:31Z","timestamp":1371077011000},"page":"3008-3017","source":"Crossref","is-referenced-by-count":49,"title":["Syndecan 4 interacts genetically with Vangl2 to regulate neural tube closure and planar cell polarity"],"prefix":"10.1242","volume":"140","author":[{"given":"Noelia","family":"Escobedo","sequence":"first","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Osvaldo","family":"Contreras","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosana","family":"Mu\u00f1oz","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marjorie","family":"Far\u00edas","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H\u00e9ctor","family":"Carrasco","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charlotte","family":"Hill","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Uyen","family":"Tran","sequence":"additional","affiliation":[{"name":"Cleveland Clinic Foundation, Lerner Research Institute, Department of Cell Biology, 9500 Euclid Avenue\/NC10, Cleveland, OH 44195, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophie E.","family":"Pryor","sequence":"additional","affiliation":[{"name":"Neural Development Unit, Institute of Child Health, University College London, 30 Guilford Street, London WC1N 1EH, UK."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oliver","family":"Wessely","sequence":"additional","affiliation":[{"name":"Cleveland Clinic Foundation, Lerner Research Institute, Department of Cell Biology, 9500 Euclid Avenue\/NC10, Cleveland, OH 44195, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew J.","family":"Copp","sequence":"additional","affiliation":[{"name":"Neural Development Unit, Institute of Child Health, University College London, 30 Guilford Street, London WC1N 1EH, UK."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan","family":"Larra\u00edn","sequence":"additional","affiliation":[{"name":"Center for Aging and Regeneration, Millennium Nucleus in Regenerative Biology, Faculty of Biological Sciences, P. Universidad Cat\u00f3lica de Chile, Alameda 340 Santiago, Chile."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","reference":[{"key":"2021042607201247800_R1","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1016\/j.devcel.2011.08.007","article-title":"A syndecan-4 hair trigger initiates wound healing through caveolin- and RhoG-regulated integrin endocytosis","volume":"21","author":"Bass","year":"2011","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R2","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/j.stem.2012.09.015","article-title":"Fibronectin regulates Wnt7a signaling and satellite cell expansion","volume":"12","author":"Bentzinger","year":"2013","journal-title":"Cell Stem Cell"},{"key":"2021042607201247800_R3","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1242\/dev.113.2.671","article-title":"Curvature of the caudal region is responsible for failure of neural tube closure in the curly tail (ct) mouse embryo","volume":"113","author":"Brook","year":"1991","journal-title":"Development"},{"key":"2021042607201247800_R4","doi-asserted-by":"crossref","first-page":"1536","DOI":"10.1093\/hmg\/ddr031","article-title":"Over-expression of Grhl2 causes spina bifida in the Axial defects mutant mouse","volume":"20","author":"Brouns","year":"2011","journal-title":"Hum. Mol. Genet."},{"key":"2021042607201247800_R5","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.devcel.2010.06.008","article-title":"Epidermal wound repair is regulated by the planar cell polarity signaling pathway","volume":"19","author":"Caddy","year":"2010","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R6","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1002\/jcb.20586","article-title":"Heparan sulfate proteoglycans exert positive and negative effects in Shh activity","volume":"96","author":"Carrasco","year":"2005","journal-title":"J. Cell. Biochem."},{"key":"2021042607201247800_R7","doi-asserted-by":"crossref","first-page":"29546","DOI":"10.1074\/jbc.M110.155812","article-title":"Non-canonical Wnt signaling induces ubiquitination and degradation of Syndecan4","volume":"285","author":"Carvallo","year":"2010","journal-title":"J. Biol. Chem."},{"key":"2021042607201247800_R8","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1242\/dev.104.2.285","article-title":"A cell-type-specific abnormality of cell proliferation in mutant (curly tail) mouse embryos developing spinal neural tube defects","volume":"104","author":"Copp","year":"1988","journal-title":"Development"},{"key":"2021042607201247800_R9","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1006\/dbio.1994.1230","article-title":"Developmental basis of severe neural tube defects in the loop-tail (Lp) mutant mouse: use of microsatellite DNA markers to identify embryonic genotype","volume":"165","author":"Copp","year":"1994","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R10","first-page":"135","article-title":"Neurulation and neural tube closure defects","volume":"136","author":"Copp","year":"2000","journal-title":"Methods Mol. Biol."},{"key":"2021042607201247800_R11","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1038\/nrg1181","article-title":"The genetic basis of mammalian neurulation","volume":"4","author":"Copp","year":"2003","journal-title":"Nat. Rev. Genet."},{"key":"2021042607201247800_R12","doi-asserted-by":"crossref","first-page":"2231","DOI":"10.1101\/gad.1214204","article-title":"Essential and separable roles for Syndecan-3 and Syndecan-4 in skeletal muscle development and regeneration","volume":"18","author":"Cornelison","year":"2004","journal-title":"Genes Dev."},{"key":"2021042607201247800_R13","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1146\/annurev-cellbio-100109-104126","article-title":"Transmembrane signaling proteoglycans","volume":"26","author":"Couchman","year":"2010","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"2021042607201247800_R14","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1038\/ncb1784","article-title":"Planar polarization in embryonic epidermis orchestrates global asymmetric morphogenesis of hair follicles","volume":"10","author":"Devenport","year":"2008","journal-title":"Nat. Cell Biol."},{"key":"2021042607201247800_R15","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1002\/ajmg.a.30766","article-title":"Analysis of the planar cell polarity gene Vangl2 and its co-expressed paralogue Vangl1 in neural tube defect patients","volume":"136","author":"Doudney","year":"2005","journal-title":"Am. J. Med. Genet. A."},{"key":"2021042607201247800_R16","doi-asserted-by":"crossref","first-page":"R9","DOI":"10.1172\/JCI10559","article-title":"Delayed wound repair and impaired angiogenesis in mice lacking syndecan-4","volume":"107","author":"Echtermeyer","year":"2001","journal-title":"J. Clin. Invest."},{"key":"2021042607201247800_R17","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1111\/1523-1747.ep12365571","article-title":"Syndecans-1 and -4 are induced during wound repair of neonatal but not fetal skin","volume":"107","author":"Gallo","year":"1996","journal-title":"J. Invest. Dermatol."},{"key":"2021042607201247800_R18","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1016\/j.devcel.2011.01.001","article-title":"Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2","volume":"20","author":"Gao","year":"2011","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R19","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1006\/dbio.2002.0673","article-title":"The planar cell polarity gene strabismus regulates convergence and extension and neural fold closure in Xenopus","volume":"247","author":"Goto","year":"2002","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R20","doi-asserted-by":"crossref","first-page":"3445","DOI":"10.1021\/bi902180m","article-title":"Molecular and cellular mechanisms underlying neural tube defects in the loop-tail mutant mouse","volume":"49","author":"Gravel","year":"2010","journal-title":"Biochemistry"},{"key":"2021042607201247800_R21","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.devcel.2011.06.011","article-title":"Planar cell polarity: coordinating morphogenetic cell behaviors with embryonic polarity","volume":"21","author":"Gray","year":"2011","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R22","doi-asserted-by":"crossref","first-page":"2640","DOI":"10.1093\/hmg\/ddm221","article-title":"Increased expression of Grainyhead-like-3 rescues spina bifida in a folate-resistant mouse model","volume":"16","author":"Gustavsson","year":"2007","journal-title":"Hum. Mol. Genet."},{"key":"2021042607201247800_R23","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1002\/dvdy.22577","article-title":"A novel hypomorphic Looptail allele at the planar cell polarity Vangl2 gene","volume":"240","author":"Guyot","year":"2011","journal-title":"Dev. Dyn."},{"key":"2021042607201247800_R24","doi-asserted-by":"crossref","first-page":"795","DOI":"10.1021\/bi101286d","article-title":"Loss of membrane targeting of Vangl proteins causes neural tube defects","volume":"50","author":"Iliescu","year":"2011","journal-title":"Biochemistry"},{"key":"2021042607201247800_R25","doi-asserted-by":"crossref","first-page":"5249","DOI":"10.1074\/jbc.275.8.5249","article-title":"Syndecan-4 deficiency impairs focal adhesion formation only under restricted conditions","volume":"275","author":"Ishiguro","year":"2000","journal-title":"J. Biol. Chem."},{"key":"2021042607201247800_R26","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/S0070-2153(08)00808-9","article-title":"Primary cilia in planar cell polarity regulation of the inner ear","volume":"85","author":"Jones","year":"2008","journal-title":"Curr. Top. Dev. Biol."},{"key":"2021042607201247800_R27","doi-asserted-by":"crossref","first-page":"1432","DOI":"10.1056\/NEJMoa060651","article-title":"Mutations in VANGL1 associated with neural-tube defects","volume":"356","author":"Kibar","year":"2007","journal-title":"N. Engl. J. Med."},{"key":"2021042607201247800_R28","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1016\/j.stem.2009.03.013","article-title":"Wnt7a activates the planar cell polarity pathway to drive the symmetric expansion of satellite stem cells","volume":"4","author":"Le Grand","year":"2009","journal-title":"Cell Stem Cell"},{"key":"2021042607201247800_R29","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1038\/nature02677","article-title":"PTK7\/CCK-4 is a novel regulator of planar cell polarity in vertebrates","volume":"430","author":"Lu","year":"2004","journal-title":"Nature"},{"key":"2021042607201247800_R30","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1006\/dbio.1998.9032","article-title":"Functional interactions of genes mediating convergent extension, knypek and trilobite, during the partitioning of the eye primordium in zebrafish","volume":"203","author":"Marlow","year":"1998","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R31","doi-asserted-by":"crossref","first-page":"1771","DOI":"10.1242\/dev.017350","article-title":"Directional migration of neural crest cells in vivo is regulated by Syndecan-4\/Rac1 and non-canonical Wnt signaling\/RhoA","volume":"135","author":"Matthews","year":"2008","journal-title":"Development"},{"key":"2021042607201247800_R32","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1038\/ncb2002","article-title":"Sec24b selectively sorts Vangl2 to regulate planar cell polarity during neural tube closure","volume":"12","author":"Merte","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"2021042607201247800_R33","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1038\/nature01618","article-title":"Identification of Vangl2 and Scrb1 as planar polarity genes in mammals","volume":"423","author":"Montcouquiol","year":"2003","journal-title":"Nature"},{"key":"2021042607201247800_R34","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1038\/nrm2289","article-title":"Synergistic control of cell adhesion by integrins and syndecans","volume":"8","author":"Morgan","year":"2007","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2021042607201247800_R35","doi-asserted-by":"crossref","first-page":"1417","DOI":"10.1242\/dev.073239","article-title":"Cell movements of the deep layer of non-neural ectoderm underlie complete neural tube closure in Xenopus","volume":"139","author":"Morita","year":"2012","journal-title":"Development"},{"key":"2021042607201247800_R36","doi-asserted-by":"crossref","first-page":"492","DOI":"10.1038\/ncb1399","article-title":"Syndecan-4 regulates non-canonical Wnt signalling and is essential forconvergent and extension movements in Xenopus embryos","volume":"8","author":"Mu\u00f1oz","year":"2006","journal-title":"Nat. Cell Biol."},{"key":"2021042607201247800_R37","doi-asserted-by":"crossref","first-page":"2593","DOI":"10.1093\/hmg\/10.22.2593","article-title":"Severe neural tube defects in the loop-tail mouse result from mutation of Lpp1, a novel gene involved in floor plate specification","volume":"10","author":"Murdoch","year":"2001","journal-title":"Hum. Mol. Genet."},{"key":"2021042607201247800_R38","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1093\/hmg\/ddg014","article-title":"Disruption of scribble (Scrb1) causes severe neural tube defects in the circletail mouse","volume":"12","author":"Murdoch","year":"2003","journal-title":"Hum. Mol. Genet."},{"key":"2021042607201247800_R39","doi-asserted-by":"crossref","first-page":"e18410","DOI":"10.1371\/journal.pone.0018410","article-title":"Delayed re-epithelialization in periostin-deficient mice during cutaneous wound healing","volume":"6","author":"Nishiyama","year":"2011","journal-title":"PLoS ONE"},{"key":"2021042607201247800_R40","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1016\/j.ydbio.2010.06.010","article-title":"Dynamic imaging of mammalian neural tube closure","volume":"344","author":"Pyrgaki","year":"2010","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R41","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.ydbio.2011.02.027","article-title":"Grainyhead-like 2 regulates neural tube closure and adhesion molecule expression during neural fold fusion","volume":"353","author":"Pyrgaki","year":"2011","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R42","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.ydbio.2007.03.011","article-title":"Wnt5a functions in planar cell polarity regulation in mice","volume":"306","author":"Qian","year":"2007","journal-title":"Dev. Biol."},{"key":"2021042607201247800_R43","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1002\/humu.21662","article-title":"Mutations in the planar cell polarity genes CELSR1 and SCRIB are associated with the severe neural tube defect craniorachischisis","volume":"33","author":"Robinson","year":"2012","journal-title":"Hum. Mutat."},{"key":"2021042607201247800_R44","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1016\/j.modgep.2006.07.007","article-title":"Tissue, cellular and sub-cellular localization of the Vangl2 protein during embryonic development: effect of the Lp mutation","volume":"7","author":"Torban","year":"2007","journal-title":"Gene Expr. Patterns"},{"key":"2021042607201247800_R45","doi-asserted-by":"crossref","first-page":"2581","DOI":"10.1242\/dev.128.13.2581","article-title":"Xenopus Dishevelled signaling regulates both neural and mesodermal convergent extension: parallel forces elongating the body axis","volume":"128","author":"Wallingford","year":"2001","journal-title":"Development"},{"key":"2021042607201247800_R46","doi-asserted-by":"crossref","first-page":"5815","DOI":"10.1242\/dev.00123","article-title":"Neural tube closure requires Dishevelled-dependent convergent extension of the midline","volume":"129","author":"Wallingford","year":"2002","journal-title":"Development"},{"key":"2021042607201247800_R47","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1016\/S1534-5807(02)00197-1","article-title":"Convergent extension: the molecular control of polarized cell movement during embryonic development","volume":"2","author":"Wallingford","year":"2002","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R48","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1242\/dev.02772","article-title":"Tissue\/planar cell polarity in vertebrates: new insights and new questions","volume":"134","author":"Wang","year":"2007","journal-title":"Development"},{"key":"2021042607201247800_R49","doi-asserted-by":"crossref","first-page":"2141","DOI":"10.1242\/jcs.097964","article-title":"VANGL2 regulates membrane trafficking of MMP14 to control cell polarity and migration","volume":"125","author":"Williams","year":"2012","journal-title":"J. Cell Sci."},{"key":"2021042607201247800_R50","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.devcel.2008.05.007","article-title":"Cthrc1 selectively activates the planar cell polarity pathway of Wnt signaling by stabilizing the Wnt-receptor complex","volume":"15","author":"Yamamoto","year":"2008","journal-title":"Dev. Cell"},{"key":"2021042607201247800_R51","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1242\/dev.000380","article-title":"Convergent extension, planar-cell-polarity signalling and initiation of mouse neural tube closure","volume":"134","author":"Ybot-Gonzalez","year":"2007","journal-title":"Development"},{"key":"2021042607201247800_R52","doi-asserted-by":"crossref","first-page":"e31988","DOI":"10.1371\/journal.pone.0031988","article-title":"Comparison of phenotypes between different vangl2 mutants demonstrates dominant effects of the Looptail mutation during hair cell development","volume":"7","author":"Yin","year":"2012","journal-title":"PLoS ONE"},{"key":"2021042607201247800_R53","doi-asserted-by":"crossref","first-page":"2109","DOI":"10.1242\/dev.129.9.2109","article-title":"Heparan sulphate proteoglycans and spinal neurulation in the mouse embryo","volume":"129","author":"Yip","year":"2002","journal-title":"Development"}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/journals.biologists.com\/dev\/article-pdf\/140\/14\/3008\/1565852\/3008.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/dev\/article-pdf\/140\/14\/3008\/1565852\/3008.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,26]],"date-time":"2021-04-26T06:55:56Z","timestamp":1619420156000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/140\/14\/3008\/45774\/Syndecan-4-interacts-genetically-with-Vangl2-to"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2013,7,15]]},"references-count":53,"journal-issue":{"issue":"14","published-print":{"date-parts":[[2013,7,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.091173","relation":{"has-review":[{"id-type":"doi","id":"10.3410\/f.718019262.793479160","asserted-by":"object"}]},"ISSN":["1477-9129","0950-1991"],"issn-type":[{"value":"1477-9129","type":"electronic"},{"value":"0950-1991","type":"print"}],"subject":[],"published":{"date-parts":[[2013,7,15]]}}}