{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T20:03:22Z","timestamp":1781208202049,"version":"3.54.1"},"reference-count":56,"publisher":"The Company of Biologists","issue":"24","license":[{"start":{"date-parts":[[2025,12,22]],"date-time":"2025-12-22T00:00:00Z","timestamp":1766361600000},"content-version":"vor","delay-in-days":7,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["GM136236"],"award-info":[{"award-number":["GM136236"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["GM134885"],"award-info":[{"award-number":["GM134885"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["GM125091"],"award-info":[{"award-number":["GM125091"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"crossref","award":["P20GM103432"],"award-info":[{"award-number":["P20GM103432"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100008106","name":"University of Wyoming","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100008106","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,12,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Cell fusion is a fundamental process in the development of many multicellular organisms, but its precise role in gene regulation and differentiation remains largely unknown. The Caenorhabditis elegans epidermis, which comprises multiple syncytial cells in the adult, represents a powerful model for studying cell fusion in the context of animal development. The largest of these epidermal syncytia, hyp7, integrates 139 individual nuclei through processive cell fusion mediated by the fusogenic protein EFF-1. To explore the role of cell fusion in developmental progression and associated gene expression changes, we conducted transcriptomic analyses of eff-1 fusion-defective C. elegans mutants. Our RNA-seq findings showed widespread transcriptomic changes including the enrichment of epidermal genes and molecular pathways involved in epidermal function during development. Single-molecule fluorescence in situ hybridization further validated the observed altered expression of mRNA transcripts. Moreover, bioinformatic analysis suggests that fusion may play a key role in promoting developmental progression within the epidermis. Our results underscore the significance of cell\u2013cell fusion in shaping transcriptional programs during development.<\/jats:p>","DOI":"10.1242\/dev.205089","type":"journal-article","created":{"date-parts":[[2025,11,17]],"date-time":"2025-11-17T08:43:55Z","timestamp":1763369035000},"source":"Crossref","is-referenced-by-count":2,"title":["Epidermal cell fusion promotes the transition from an embryonic to a larval transcriptome in\n                    <i>C. elegans<\/i>"],"prefix":"10.1242","volume":"152","author":[{"given":"Owen H.","family":"Funk","sequence":"first","affiliation":[{"name":"University of Wyoming Department of Molecular Biology , , Laramie, WY 82071 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7853-3275","authenticated-orcid":false,"given":"Daniel L.","family":"Levy","sequence":"additional","affiliation":[{"name":"University of Wyoming Department of Molecular Biology , , Laramie, WY 82071 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7599-4017","authenticated-orcid":false,"given":"David S.","family":"Fay","sequence":"additional","affiliation":[{"name":"University of Wyoming Department of Molecular Biology , , Laramie, WY 82071 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2025,12,22]]},"reference":[{"key":"2025122208455809400_DEV205089C1","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1242\/dev.068353","article-title":"Myoblast fusion: lessons from flies and mice","volume":"139","author":"Abmayr","year":"2012","journal-title":"Development"},{"key":"2025122208455809400_DEV205089C2","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1016\/j.tig.2013.01.011","article-title":"Genetic basis of cell-cell fusion mechanisms","volume":"29","author":"Aguilar","year":"2013","journal-title":"Trends Genet."},{"key":"2025122208455809400_DEV205089C3","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1007\/978-1-59745-250-2_4","article-title":"Cell fusion in Caenorhabditis elegans","volume":"475","author":"Alper","year":"2008","journal-title":"Methods Mol. Biol."},{"key":"2025122208455809400_DEV205089C4","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","article-title":"HTSeq--a Python framework to work with high-throughput sequencing data","volume":"31","author":"Anders","year":"2015","journal-title":"Bioinformatics"},{"key":"2025122208455809400_DEV205089C5","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1101\/gr.202663.115","article-title":"The time-resolved transcriptome of C. elegans","volume":"26","author":"Boeck","year":"2016","journal-title":"Genome Res."},{"key":"2025122208455809400_DEV205089C6","doi-asserted-by":"publisher","first-page":"e1002200","DOI":"10.1371\/journal.pgen.1002200","article-title":"The Caenorhabditis elegans GATA factor ELT-1 works through the cell proliferation regulator BRO-1 and the Fusogen EFF-1 to maintain the seam stem-like fate","volume":"7","author":"Brabin","year":"2011","journal-title":"PLoS Genet."},{"key":"2025122208455809400_DEV205089C7","doi-asserted-by":"publisher","first-page":"dev200984","DOI":"10.1242\/dev.200984","article-title":"The GATA factor ELT-3 specifies endoderm in Caenorhabditis angaria in an ancestral gene network","volume":"149","author":"Broitman-Maduro","year":"2022","journal-title":"Development"},{"key":"2025122208455809400_DEV205089C8","doi-asserted-by":"publisher","first-page":"1145","DOI":"10.1016\/j.ajpath.2019.02.006","article-title":"An overview of the derivation and function of multinucleated giant cells and their role in pathologic processes","volume":"189","author":"Brooks","year":"2019","journal-title":"Am. J. Pathol."},{"key":"2025122208455809400_DEV205089C9","doi-asserted-by":"publisher","first-page":"657","DOI":"10.1530\/REP-15-0544","article-title":"Studies of the dynamics of nuclear clustering in human syncytiotrophoblast","volume":"151","author":"Calvert","year":"2016","journal-title":"Reproduction"},{"key":"2025122208455809400_DEV205089C10","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1126\/science.aam8940","article-title":"Comprehensive single-cell transcriptional profiling of a multicellular organism","volume":"357","author":"Cao","year":"2017","journal-title":"Science"},{"key":"2025122208455809400_DEV205089C11","doi-asserted-by":"publisher","first-page":"250","DOI":"10.1055\/s-2008-1062269","article-title":"Inherited myopathies and muscular dystrophies","volume":"28","author":"Cardamone","year":"2008","journal-title":"Semin. Neurol."},{"key":"2025122208455809400_DEV205089C12","doi-asserted-by":"publisher","first-page":"1415","DOI":"10.1534\/g3.112.003830","article-title":"Large-scale screening for targeted knockouts in the Caenorhabditis elegans genome","volume":"2","author":"C. elegans Deletion Mutant Consortium","year":"2012","journal-title":"G3"},{"key":"2025122208455809400_DEV205089C13","doi-asserted-by":"publisher","first-page":"2181","DOI":"10.1016\/j.febslet.2007.03.033","article-title":"Cell-cell fusion","volume":"581","author":"Chen","year":"2007","journal-title":"FEBS Lett."},{"key":"2025122208455809400_DEV205089C14","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1002\/wdev.79","article-title":"The Caenorhabditis elegans epidermis as a model skin. I: development, patterning, and growth","volume":"1","author":"Chisholm","year":"2012","journal-title":"Wiley Interdiscip. Rev. Dev. Biol."},{"key":"2025122208455809400_DEV205089C15","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1006\/dbio.1997.8530","article-title":"The role of actin filaments in patterning the Caenorhabditis elegans cuticle","volume":"184","author":"Costa","year":"1997","journal-title":"Dev. Biol."},{"key":"2025122208455809400_DEV205089C16","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1016\/j.semcancer.2021.03.005","article-title":"Cell-cell fusions and cell-in-cell phenomena in healthy cells and cancer: lessons from protists and invertebrates","volume":"81","author":"Demin","year":"2022","journal-title":"Semin. Cancer Biol."},{"key":"2025122208455809400_DEV205089C17","doi-asserted-by":"publisher","first-page":"e202007209","DOI":"10.1083\/jcb.202007209","article-title":"Astral microtubule cross-linking safeguards uniform nuclear distribution in the Drosophila syncytium","volume":"221","author":"Deshpande","year":"2022","journal-title":"J. Cell Biol."},{"key":"2025122208455809400_DEV205089C18","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: ultrafast universal RNA-seq aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"key":"2025122208455809400_DEV205089C19","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1038\/s41580-021-00389-z","article-title":"Molecular and cellular basis of genetically inherited skeletal muscle disorders","volume":"22","author":"Dowling","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2025122208455809400_DEV205089C20","doi-asserted-by":"publisher","first-page":"1775","DOI":"10.1126\/science.1196914","article-title":"Integrative analysis of the Caenorhabditis elegans genome by the modENCODE project","volume":"330","author":"Gerstein","year":"2010","journal-title":"Science"},{"key":"2025122208455809400_DEV205089C21","doi-asserted-by":"publisher","first-page":"100748","DOI":"10.1016\/j.xpro.2021.100748","article-title":"Microinjection for precision genome editing in Caenorhabditis elegans","volume":"2","author":"Ghanta","year":"2021","journal-title":"STAR Protoc."},{"key":"2025122208455809400_DEV205089C22","doi-asserted-by":"publisher","first-page":"2533","DOI":"10.1128\/MCB.21.7.2533-2544.2001","article-title":"Activation of hypodermal differentiation in the Caenorhabditis elegans embryo by GATA transcription factors ELT-1 and ELT-3","volume":"21","author":"Gilleard","year":"2001","journal-title":"Mol. Cell. Biol."},{"key":"2025122208455809400_DEV205089C23","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1006\/dbio.1999.9202","article-title":"ELT-3: a Caenorhabditis elegans GATA factor expressed in the embryonic epidermis during morphogenesis","volume":"208","author":"Gilleard","year":"1999","journal-title":"Dev. Biol."},{"key":"2025122208455809400_DEV205089C24","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1016\/j.molcel.2013.12.013","article-title":"Extensive oscillatory gene expression during C. elegans larval development","volume":"53","author":"Hendriks","year":"2014","journal-title":"Mol. Cell"},{"key":"2025122208455809400_DEV205089C25","doi-asserted-by":"publisher","first-page":"e1234659","DOI":"10.1080\/21624054.2016.1234659","article-title":"GExplore 1.4: an expanded web interface for queries on","volume":"5","author":"Hutter","year":"2016","journal-title":"Worm"},{"key":"2025122208455809400_DEV205089C26","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/bs.ctdb.2020.12.013","article-title":"Programmed cell fusion in development and homeostasis","volume":"144","author":"Iosilevskii","year":"2021","journal-title":"Curr. Top. Dev. Biol."},{"key":"2025122208455809400_DEV205089C27","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1016\/S0168-9525(99)01857-0","article-title":"Cuticle collagen genes. Expression in Caenorhabditis elegans","volume":"16","author":"Johnstone","year":"2000","journal-title":"Trends Genet."},{"key":"2025122208455809400_DEV205089C28","doi-asserted-by":"publisher","first-page":"e1007559","DOI":"10.1371\/journal.pgen.1007559","article-title":"Transcriptome analysis of adult Caenorhabditis elegans cells reveals tissue-specific gene and isoform expression","volume":"14","author":"Kaletsky","year":"2018","journal-title":"PLoS Genet."},{"key":"2025122208455809400_DEV205089C29","doi-asserted-by":"publisher","first-page":"6375","DOI":"10.1038\/s41467-020-20064-9","article-title":"Single-nucleus transcriptomics reveals functional compartmentalization in syncytial skeletal muscle cells","volume":"11","author":"Kim","year":"2020","journal-title":"Nat. Commun."},{"key":"2025122208455809400_DEV205089C30","doi-asserted-by":"publisher","first-page":"9787","DOI":"10.1038\/s41598-021-88500-4","article-title":"A role for the fusogen eff-1 in epidermal stem cell number robustness in Caenorhabditis elegans","volume":"11","author":"Koneru","year":"2021","journal-title":"Sci. Rep."},{"key":"2025122208455809400_DEV205089C31","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1007\/s00418-008-0411-1","article-title":"Cell fusions in mammals","volume":"129","author":"Larsson","year":"2008","journal-title":"Histochem. Cell Biol."},{"key":"2025122208455809400_DEV205089C32","doi-asserted-by":"publisher","first-page":"623","DOI":"10.1016\/j.cell.2009.08.044","article-title":"Analysis of cell fate from single-cell gene expression profiles in C. elegans","volume":"139","author":"Liu","year":"2009","journal-title":"Cell"},{"key":"2025122208455809400_DEV205089C33","doi-asserted-by":"publisher","first-page":"925","DOI":"10.1016\/j.neuron.2011.12.037","article-title":"CYSL-1 interacts with the O2-sensing hydroxylase EGL-9 to promote H2S-modulated hypoxia-induced behavioral plasticity in C. elegans","volume":"73","author":"Ma","year":"2012","journal-title":"Neuron"},{"key":"2025122208455809400_DEV205089C34","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1242\/dev.075135","article-title":"Extracellular leucine-rich repeat proteins are required to organize the apical extracellular matrix and maintain epithelial junction integrity in C. elegans","volume":"139","author":"Mancuso","year":"2012","journal-title":"Development"},{"key":"2025122208455809400_DEV205089C35","doi-asserted-by":"publisher","first-page":"e9498","DOI":"10.15252\/msb.20209498","article-title":"Developmental function and state transitions of a gene expression oscillator in Caenorhabditis elegans","volume":"16","author":"Meeuse","year":"2020","journal-title":"Mol. Syst. Biol."},{"key":"2025122208455809400_DEV205089C36","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/S1534-5807(02)00129-6","article-title":"The type I membrane protein EFF-1 is essential for developmental cell fusion","volume":"2","author":"Mohler","year":"2002","journal-title":"Dev. Cell"},{"key":"2025122208455809400_DEV205089C37","doi-asserted-by":"publisher","first-page":"703","DOI":"10.1038\/nmeth.1228","article-title":"Automated analysis of embryonic gene expression with cellular resolution in C. elegans","volume":"5","author":"Murray","year":"2008","journal-title":"Nat. Methods"},{"key":"2025122208455809400_DEV205089C38","doi-asserted-by":"publisher","first-page":"567","DOI":"10.1038\/nrm1678","article-title":"Biological implications of cell fusion","volume":"6","author":"Ogle","year":"2005","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2025122208455809400_DEV205089C39","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1016\/j.tig.2015.03.011","article-title":"When bigger is better: the role of polyploidy in organogenesis","volume":"31","author":"Orr-Weaver","year":"2015","journal-title":"Trends Genet."},{"key":"2025122208455809400_DEV205089C40","doi-asserted-by":"publisher","first-page":"eaax1971","DOI":"10.1126\/science.aax1971","article-title":"A lineage-resolved molecular atlas of C. elegans embryogenesis at single-cell resolution","volume":"365","author":"Packer","year":"2019","journal-title":"Science"},{"key":"2025122208455809400_DEV205089C41","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1895\/wormbook.1.138.1","article-title":"The cuticle","volume-title":"WormBook","author":"Page","year":"2007"},{"key":"2025122208455809400_DEV205089C42","doi-asserted-by":"publisher","first-page":"e299","DOI":"10.1002\/cpz1.299","article-title":"Improved methods for single-Mmolecule fluorescence in situ hybridization and immunofluorescence in Caenorhabditis elegans embryos","volume":"1","author":"Parker","year":"2021","journal-title":"Curr. Protoc."},{"key":"2025122208455809400_DEV205089C43","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1038\/nmeth.4197","article-title":"Salmon provides fast and bias-aware quantification of transcript expression","volume":"14","author":"Patro","year":"2017","journal-title":"Nat. Methods"},{"key":"2025122208455809400_DEV205089C44","doi-asserted-by":"publisher","first-page":"6374","DOI":"10.1038\/s41467-020-20063-w","article-title":"Single-nucleus RNA-seq identifies transcriptional heterogeneity in multinucleated skeletal myofibers","volume":"11","author":"Petrany","year":"2020","journal-title":"Nat. Commun."},{"key":"2025122208455809400_DEV205089C45","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1006\/dbio.1994.1041","article-title":"Cell fusions in the developing epithelial of C. elegans","volume":"161","author":"Podbilewicz","year":"1994","journal-title":"Dev. Biol."},{"key":"2025122208455809400_DEV205089C46","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1016\/j.devcel.2006.09.004","article-title":"The C. elegans developmental fusogen EFF-1 mediates homotypic fusion in heterologous cells and in vivo","volume":"11","author":"Podbilewicz","year":"2006","journal-title":"Dev. Cell"},{"key":"2025122208455809400_DEV205089C47","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/0012-1606(86)90358-1","article-title":"Caenorhabditis elegans morphogenesis: the role of the cytoskeleton in elongation of the embryo","volume":"117","author":"Priess","year":"1986","journal-title":"Dev. Biol."},{"key":"2025122208455809400_DEV205089C48","doi-asserted-by":"publisher","first-page":"13500","DOI":"10.1093\/nar\/gku1228","article-title":"Prediction of DNA binding motifs from 3D models of transcription factors; identifying TLX3 regulated genes","volume":"42","author":"Pujato","year":"2014","journal-title":"Nucleic Acids Res."},{"key":"2025122208455809400_DEV205089C49","doi-asserted-by":"publisher","first-page":"W160","DOI":"10.1093\/nar\/gkw257","article-title":"deepTools2: a next generation web server for deep-sequencing data analysis","volume":"44","author":"Ram\u00edrez","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2025122208455809400_DEV205089C50","doi-asserted-by":"publisher","first-page":"R786","DOI":"10.1016\/j.cub.2011.06.042","article-title":"Nuclear and genome dynamics in multinucleate ascomycete fungi","volume":"21","author":"Roper","year":"2011","journal-title":"Curr. Biol."},{"key":"2025122208455809400_DEV205089C51","doi-asserted-by":"publisher","first-page":"1587","DOI":"10.1016\/j.cub.2004.07.059","article-title":"EFF-1 is sufficient to initiate and execute tissue-specific cell fusion in C. elegans","volume":"14","author":"Shemer","year":"2004","journal-title":"Curr. Biol."},{"key":"2025122208455809400_DEV205089C52","doi-asserted-by":"publisher","first-page":"1205","DOI":"10.1534\/genetics.106.063594","article-title":"The Caenorhabditis elegans rhy-1 gene inhibits HIF-1 hypoxia-inducible factor activity in a negative feedback loop that does not include vhl-1","volume":"174","author":"Shen","year":"2006","journal-title":"Genetics"},{"key":"2025122208455809400_DEV205089C53","doi-asserted-by":"publisher","first-page":"e0146874","DOI":"10.1371\/journal.pone.0146874","article-title":"The EFF-1A cytoplasmic domain influences hypodermal cell fusions in C. elegans but is not dependent on 14-3-3 proteins","volume":"11","author":"Shinn-Thomas","year":"2016","journal-title":"PLoS ONE"},{"key":"2025122208455809400_DEV205089C54","doi-asserted-by":"publisher","first-page":"iyae072","DOI":"10.1093\/genetics\/iyae072","article-title":"The Caenorhabditis elegans cuticle and precuticle: a model for studying dynamic apical extracellular matrices in vivo","volume":"227","author":"Sundaram","year":"2024","journal-title":"Genetics"},{"key":"2025122208455809400_DEV205089C55","doi-asserted-by":"publisher","first-page":"e165","DOI":"10.1093\/nar\/gkw784","article-title":"smiFISH and FISH-quant - a flexible single RNA detection approach with super-resolution capability","volume":"44","author":"Tsanov","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2025122208455809400_DEV205089C56","doi-asserted-by":"publisher","first-page":"835099","DOI":"10.3389\/fgene.2022.835099","article-title":"Heterogeneous skeletal muscle cell and nucleus populations identified by single-cell and single-nucleus resolution transcriptome assays","volume":"13","author":"Williams","year":"2022","journal-title":"Front. Genet."}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205089\/3673576\/dev205089.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205089\/3678314\/dev205089.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205089\/3678314\/dev205089.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,22]],"date-time":"2025-12-22T13:46:04Z","timestamp":1766411164000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/152\/24\/dev205089\/370195\/Epidermal-cell-fusion-promotes-the-transition-from"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,15]]},"references-count":56,"journal-issue":{"issue":"24","published-print":{"date-parts":[[2025,12,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.205089","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"value":"0950-1991","type":"print"},{"value":"1477-9129","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,15]]},"article-number":"dev205089"}}