{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T19:23:23Z","timestamp":1790191403899,"version":"4.1.0"},"reference-count":35,"publisher":"The Company of Biologists","issue":"21","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Fonds de Recherche du Qu\u00e9bec - Sant\u00e9","award":["296557"],"award-info":[{"award-number":["296557"]}],"id":[{"id":"https:\/\/ror.org\/02eqrsj93","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Fonds de Recherche du Qu\u00e9bec - Sant\u00e9","award":["360833"],"award-info":[{"award-number":["360833"]}],"id":[{"id":"https:\/\/ror.org\/02eqrsj93","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Fonds de Recherche du Qu\u00e9bec - Sant\u00e9","award":["2002751"],"award-info":[{"award-number":["2002751"]}],"id":[{"id":"https:\/\/ror.org\/02eqrsj93","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Natural Sciences and Engineering Research Council of Canada","id":[{"id":"https:\/\/ror.org\/01h531d29","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Canada Graduate Scholarships","award":["202211FBD-493573-92422"],"award-info":[{"award-number":["202211FBD-493573-92422"]}]},{"name":"Canadian Institutes of Health Research","award":["360833"],"award-info":[{"award-number":["360833"]}],"id":[{"id":"https:\/\/ror.org\/01gavpb45","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Cancer Research Society","award":["1502616"],"award-info":[{"award-number":["1502616"]}],"id":[{"id":"https:\/\/ror.org\/00t38a349","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Cancer Research Society","award":["2002751"],"award-info":[{"award-number":["2002751"]}],"id":[{"id":"https:\/\/ror.org\/00t38a349","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Universit\u00e9 Laval","id":[{"id":"https:\/\/ror.org\/04sjchr03","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,11,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>The epidermis is a stratified epithelium in which keratinocytes differentiate as they move from basal to suprabasal layers, a process during which tissue integrity relies on the coordinated remodeling of intercellular junctions and the cytoskeleton. Unlike simple epithelia, which form a continuous zonula adherens, suprabasal cells in the epidermis lack a canonical junction belt, with E-cadherin-based junctions distributed across the entire cell surface, highlighting distinct regulatory mechanisms that remain poorly understood. Here, we identify the CDC42 guanine nucleotide exchange factor FGD1, mutated in Aarskog\u2013Scott syndrome, as a regulator of suprabasal junction architecture. Using stratified human keratinocytes and quantitative imaging, we show that loss of FGD1 induces elongated radial actin bundles and tortuous, tension-associated adherens junctions resembling the stress fiber-anchored discontinuous (focal) adherens junctions described in endothelial cells. These defects are accompanied by altered tissue mechanics. Inhibition of ROCK or myosin II restores junction organization, demonstrating that FGD1 restrains actomyosin contractility to promote suprabasal junction maturation. Several disease-associated FGD1 variants fail to rescue these defects. Together, our findings identify FGD1 as a regulator of suprabasal junction maturation.<\/jats:p>","DOI":"10.1242\/jcs.264937","type":"journal-article","created":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T09:43:19Z","timestamp":1786095799000},"source":"Crossref","is-referenced-by-count":1,"title":["FGD1 restrains actomyosin contractility to promote suprabasal junction maturation in stratified keratinocytes"],"prefix":"10.1242","volume":"139","author":[{"given":"Diana I.","family":"Hotea","sequence":"first","affiliation":[{"name":"Universit\u00e9 Laval, Qu\u00e9bec 1 D\u00e9partement de biologie mol\u00e9culaire, biochimie m\u00e9dicale et pathologie, Facult\u00e9 de m\u00e9decine , , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexandra","family":"Jett\u00e9","sequence":"additional","affiliation":[{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy","family":"Beaudin","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Laval, Qu\u00e9bec 1 D\u00e9partement de biologie mol\u00e9culaire, biochimie m\u00e9dicale et pathologie, Facult\u00e9 de m\u00e9decine , , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mait\u00e9","family":"Lauzon","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Laval, Qu\u00e9bec 1 D\u00e9partement de biologie mol\u00e9culaire, biochimie m\u00e9dicale et pathologie, Facult\u00e9 de m\u00e9decine , , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Krithika","family":"Kalyanakrishnan","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Laval, Qu\u00e9bec 1 D\u00e9partement de biologie mol\u00e9culaire, biochimie m\u00e9dicale et pathologie, Facult\u00e9 de m\u00e9decine , , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7198-3257","authenticated-orcid":false,"given":"M\u00e9lanie","family":"Laurin","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Laval, Qu\u00e9bec 1 D\u00e9partement de biologie mol\u00e9culaire, biochimie m\u00e9dicale et pathologie, Facult\u00e9 de m\u00e9decine , , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Oncology Research Axis, CHU de Qu\u00e9bec - Universit\u00e9 Laval Research Center, Qu\u00e9bec 2 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche sur le cancer de l'Universit\u00e9 Laval (CRC), Qu\u00e9bec 3 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]},{"name":"Centre de recherche en organog\u00e9n\u00e8se exp\u00e9rimentale (LOEX), Qu\u00e9bec 4 , Qu\u00e9bec G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"2026092303324746500_JCS264937C1","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.yexcr.2017.03.053","article-title":"Rho GTPases and actomyosin: partners in regulating epithelial cell-cell junction structure and function","volume":"358","author":"Arnold","year":"2017","journal-title":"Exp. Cell Res."},{"key":"2026092303324746500_JCS264937C2","doi-asserted-by":"publisher","first-page":"1421","DOI":"10.1083\/jcb.137.6.1421","article-title":"The small GTPases Rho and Rac are required for the establishment of cadherin-dependent cell-cell contacts","volume":"137","author":"Braga","year":"1997","journal-title":"J. Cell Biol."},{"key":"2026092303324746500_JCS264937C3","doi-asserted-by":"publisher","first-page":"1449","DOI":"10.1083\/jcb.141.6.1449","article-title":"Tyrosine phosphorylation and src family kinases control keratinocyte cell-cell adhesion","volume":"141","author":"Calautti","year":"1998","journal-title":"J. Cell Biol."},{"key":"2026092303324746500_JCS264937C4","doi-asserted-by":"publisher","first-page":"e202302196","DOI":"10.26508\/lsa.202302196","article-title":"MAP4K4 regulates forces at cell\u2013cell and cell\u2013matrix adhesions to promote collective cell migration","volume":"6","author":"Delsin","year":"2023","journal-title":"Life Sci. Alliance"},{"key":"2026092303324746500_JCS264937C5","doi-asserted-by":"publisher","first-page":"1480","DOI":"10.1016\/j.yexcr.2008.11.012","article-title":"Cdc42 is crucial for the maturation of primordial cell junctions in keratinocytes independent of Rac1","volume":"315","author":"Du","year":"2009","journal-title":"Exp. Cell Res."},{"key":"2026092303324746500_JCS264937C6","doi-asserted-by":"publisher","first-page":"1827","DOI":"10.1083\/jcb.201708103","article-title":"Branched actin networks push against each other at adherens junctions to maintain cell-cell adhesion","volume":"217","author":"Efimova","year":"2018","journal-title":"J. Cell Biol."},{"key":"2026092303324746500_JCS264937C7","doi-asserted-by":"publisher","first-page":"7","DOI":"10.3389\/fcell.2019.00007","article-title":"Identification of CDC42 effectors operating in FGD1-dependent trafficking at the Golgi","volume":"7","author":"Egorov","year":"2019","journal-title":"Front. Cell Dev. Biol."},{"key":"2026092303324746500_JCS264937C8","doi-asserted-by":"publisher","first-page":"1537","DOI":"10.1016\/S0002-9440(10)65025-0","article-title":"Genetic and epigenetic changes in human epithelial cells immortalized by telomerase","volume":"156","author":"Farwell","year":"2000","journal-title":"Am. J. Pathol."},{"key":"2026092303324746500_JCS264937C9","doi-asserted-by":"publisher","first-page":"1981","DOI":"10.1093\/hmg\/ddg209","article-title":"Fgd1, the Cdc42 GEF responsible for Faciogenital Dysplasia, directly interacts with cortactin and mAbp1 to modulate cell shape","volume":"12","author":"Hou","year":"2003","journal-title":"Hum. Mol. Genet."},{"key":"2026092303324746500_JCS264937C10","doi-asserted-by":"publisher","first-page":"a040840","DOI":"10.1101\/cshperspect.a040840","article-title":"Building and maintaining the skin","volume":"14","author":"Hsu","year":"2022","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"2026092303324746500_JCS264937C11","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1083\/jcb.201108120","article-title":"Vinculin associates with endothelial VE-cadherin junctions to control force-dependent remodeling","volume":"196","author":"Huveneers","year":"2012","journal-title":"J. Cell Biol."},{"key":"2026092303324746500_JCS264937C12","doi-asserted-by":"publisher","first-page":"jcs263938","DOI":"10.1242\/jcs.263938","article-title":"Piezo1 balances membrane and cortex tension to stabilize intercellular junctions and maintain the epithelial barrier","volume":"138","author":"Javed","year":"2025","journal-title":"J. Cell Sci."},{"key":"2026092303324746500_JCS264937C13","doi-asserted-by":"publisher","first-page":"2422","DOI":"10.1021\/bi036173t","article-title":"Effect of Fgd1 on cortactin in Arp2\/3 complex-mediated actin assembly","volume":"43","author":"Kim","year":"2004","journal-title":"Biochemistry"},{"key":"2026092303324746500_JCS264937C35","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1038\/s41597-022-01654-2","article-title":"RSDB: A rare skin disease database to link drugs with potential drug targets for rare skin diseases","volume":"9","author":"Kuo","year":"2022","journal-title":"Sci. Data"},{"key":"2026092303324746500_JCS264937C14","doi-asserted-by":"publisher","first-page":"e50226","DOI":"10.7554\/eLife.50226","article-title":"An RNAi screen unravels the complexities of rho GTPase networks in skin morphogenesis","volume":"8","author":"Laurin","year":"2019","journal-title":"eLife"},{"key":"2026092303324746500_JCS264937C15","doi-asserted-by":"publisher","first-page":"2257","DOI":"10.1007\/s00431-024-05484-9","article-title":"FGD1-related Aarskog\u2013Scott syndrome: Identification of four novel variations and a literature review of clinical and molecular aspects","volume":"183","author":"Li","year":"2024","journal-title":"Eur. J. Pediatr."},{"key":"2026092303324746500_JCS264937C16","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/1741-7007-8-11","article-title":"Adherens junctions connect stress fibres between adjacent endothelial cells","volume":"8","author":"Mill\u00e1n","year":"2010","journal-title":"BMC Biol."},{"key":"2026092303324746500_JCS264937C17","doi-asserted-by":"publisher","first-page":"1053","DOI":"10.1038\/s41467-018-03414-6","article-title":"Desmosomal cadherin association with Tctex-1 and cortactin-Arp2\/3 drives perijunctional actin polymerization to promote keratinocyte delamination","volume":"9","author":"Nekrasova","year":"2018","journal-title":"Nat. Commun."},{"key":"2026092303324746500_JCS264937C18","doi-asserted-by":"publisher","first-page":"a041518","DOI":"10.1101\/cshperspect.a041518","article-title":"Mechanochemical principles of epidermal tissue dynamics","volume":"17","author":"Niessen","year":"2025","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"2026092303324746500_JCS264937C19","doi-asserted-by":"publisher","first-page":"1628","DOI":"10.1016\/S0960-9822(02)70786-0","article-title":"Faciogenital dysplasia protein (FGD1) and Vav, two related proteins required for normal embryonic development, are upstream regulators of Rho GTPases","volume":"6","author":"Olson","year":"1996","journal-title":"Curr. Biol."},{"key":"2026092303324746500_JCS264937C20","doi-asserted-by":"publisher","first-page":"669","DOI":"10.1016\/0092-8674(94)90552-5","article-title":"Isolation and characterization of the faciogenital dysplasia (Aarskog-Scott Syndrome) gene: a putative Rho\/Rat guanine nucleotide exchange factor","volume":"79","author":"Pasteris","year":"1994","journal-title":"Cell"},{"key":"2026092303324746500_JCS264937C21","doi-asserted-by":"publisher","first-page":"1250","DOI":"10.1038\/s41467-017-01170-7","article-title":"E-cadherin integrates mechanotransduction and EGFR signaling to control junctional tissue polarization and tight junction positioning","volume":"8","author":"R\u00fcbsam","year":"2017","journal-title":"Nat. Commun."},{"key":"2026092303324746500_JCS264937C22","doi-asserted-by":"publisher","first-page":"7743","DOI":"10.1038\/s41598-023-33485-5","article-title":"Polarity signaling balances epithelial contractility and mechanical resistance","volume":"13","author":"R\u00fcbsam","year":"2023","journal-title":"Sci. Rep."},{"key":"2026092303324746500_JCS264937C23","doi-asserted-by":"publisher","first-page":"208","DOI":"10.3390\/vaccines10020208","article-title":"Pitfalls in the application of dispase-based keratinocyte dissociation assay for in vitro analysis of pemphigus vulgaris","volume":"10","author":"Schmidt","year":"2022","journal-title":"Vaccines (Basel)"},{"key":"2026092303324746500_JCS264937C24","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1038\/nrm3175","article-title":"Deconstructing the skin: Cytoarchitectural determinants of epidermal morphogenesis","volume":"12","author":"Simpson","year":"2011","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026092303324746500_JCS264937C25","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/j.devcel.2018.10.025","article-title":"Adherens junction length during tissue contraction is controlled by the mechanosensitive activity of actomyosin and junctional recycling","volume":"47","author":"Sumi","year":"2018","journal-title":"Dev. Cell"},{"key":"2026092303324746500_JCS264937C26","doi-asserted-by":"publisher","first-page":"383","DOI":"10.1016\/bs.ctdb.2014.11.027","article-title":"Cell adhesion in epidermal development and barrier formation","volume-title":"Curr. Top. Dev. Biol.","author":"Sumigray","year":"2015"},{"key":"2026092303324746500_JCS264937C27","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1297\/cpe.2023-0027","article-title":"Clinical Pediatric Endocrinology Mutation-in-Brief A novel missense variant of FGD1 disrupts critical cysteine residues of the FYVE domain in Japanese siblings with Aarskog-Scott syndrome","volume":"33","author":"Takahashi","year":"2024","journal-title":"Clin Pediatr Endocrinol"},{"key":"2026092303324746500_JCS264937C28","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1016\/S1534-5807(02)00259-9","article-title":"Actin cable dynamics and Rho\/Rock orchestrate a polarized cytoskeletal architercture in the early steps of assembling a stratified epithelium","volume":"3","author":"Vaezi","year":"2002","journal-title":"Dev. Cell"},{"key":"2026092303324746500_JCS264937C29","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1016\/S0092-8674(00)81559-7","article-title":"Directed actin polymerization is the driving force for epithelial cell-cell adhesion","volume":"100","author":"Vasioukhin","year":"2000","journal-title":"Cell"},{"key":"2026092303324746500_JCS264937C30","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1101\/gad.361406","article-title":"Cdc42 controls progenitor cell differentiation and \u03b2-catenin turnover in skin","volume":"20","author":"Wu","year":"2006","journal-title":"Genes Dev."},{"key":"2026092303324746500_JCS264937C31","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1038\/ncb2055","article-title":"\u03b1-Catenin as a tension transducer that induces adherens junction development","volume":"12","author":"Yonemura","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"2026092303324746500_JCS264937C32","doi-asserted-by":"publisher","first-page":"6652957","DOI":"10.1155\/2021\/6652957","article-title":"The prevalence of clinical features in patients with aarskog-scott syndrome and assessment of genotype-phenotype correlation: a systematic review","volume":"2021","author":"Zanetti Drumond","year":"2021","journal-title":"Genet. Res. (Camb)"},{"key":"2026092303324746500_JCS264937C33","doi-asserted-by":"publisher","first-page":"5065","DOI":"10.7150\/thno.34014","article-title":"Cdc42 deficiency leads to epidermal barrier dysfunction by regulating intercellular junctions and keratinization of epidermal cells during mouse skin development","volume":"9","author":"Zhang","year":"2019","journal-title":"Theranostics"},{"key":"2026092303324746500_JCS264937C34","doi-asserted-by":"publisher","first-page":"33169","DOI":"10.1074\/jbc.271.52.33169","article-title":"The faciogenital dysplasia gene product FGD1 functions as a Cdc42Hs-specific guanine-nucleotide exchange factor","volume":"271","author":"Zheng","year":"1996"}],"container-title":["Journal of Cell Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264937\/3711695\/jcs264937.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264937\/3719052\/jcs264937.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264937\/3719052\/jcs264937.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T07:32:58Z","timestamp":1790148778000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jcs\/article\/139\/21\/jcs264937\/372997\/FGD1-restrains-actomyosin-contractility-to-promote"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,23]]},"references-count":35,"journal-issue":{"issue":"21","published-print":{"date-parts":[[2026,11,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jcs.264937","relation":{},"ISSN":["0021-9533","1477-9137"],"issn-type":[{"value":"0021-9533","type":"print"},{"value":"1477-9137","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,23]]},"article-number":"jcs264937"}}