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However, the underlying mechanisms in vertebrates are poorly understood. In zebrafish, Glypican 4 (Gpc4) is required for convergence and extension (C&amp;E) of both the mesoderm and endoderm. Here, we show that transgenic expression of GFP-Gpc4 in the endoderm of gpc4 mutants rescued C&amp;E defects in all germ layers. The rescue of mesoderm was likely mediated by Wnt5b and Wnt11f2 and depended on signaling filopodia rather than on cleavage of the Gpc4 GPI anchor. Gpc4 bound both Wnt5b and Wnt11f2 and regulated formation of the filopodia that transport Wnt5b and Wnt11f2 to neighboring cells. Moreover, this rescue was suppressed by blocking signaling filopodia that extend from endodermal cells. Thus, GFP-Gpc4\u2013labeled protrusions that emanated from endodermal cells transported Wnt5b and Wnt11f2 to other germ layers, rescuing the C&amp;E defects caused by a gpc4 deficiency. Our study reveals a new mechanism that could explain in vivo morphogen distribution involving Gpc4.<\/jats:p>","DOI":"10.1083\/jcb.202009082","type":"journal-article","created":{"date-parts":[[2021,9,9]],"date-time":"2021-09-09T02:22:10Z","timestamp":1631154130000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":24,"title":["Glypican 4 mediates Wnt transport between germ layers via signaling filopodia"],"prefix":"10.1083","volume":"220","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5656-9632","authenticated-orcid":true,"given":"Bo","family":"Hu","sequence":"first","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4885-5684","authenticated-orcid":true,"given":"Juan J.","family":"Rodriguez","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anurag","family":"Kakkerla Balaraju","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanyuan","family":"Gao","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nhan T.","family":"Nguyen","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heston","family":"Steen","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9771-4274","authenticated-orcid":true,"given":"Saeb","family":"Suhaib","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Songhai","family":"Chen","sequence":"additional","affiliation":[{"name":"Department of Neuroscience and Pharmacology, Carver College of Medicine, The University of Iowa, Iowa City, IA 2"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2179-7031","authenticated-orcid":true,"given":"Fang","family":"Lin","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Cell Biology, Carver College of Medicine, The University of Iowa, Iowa City, IA 1"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2021,9,30]]},"reference":[{"key":"2023072515365372400_bib1","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1006\/dbio.1999.9441","article-title":"casanova plays an early and essential role in endoderm formation in zebrafish","volume":"215","author":"Alexander","year":"1999","journal-title":"Dev. 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