{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,22]],"date-time":"2026-03-22T08:22:25Z","timestamp":1774167745877,"version":"3.50.1"},"reference-count":107,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,6,4]],"date-time":"2021-06-04T00:00:00Z","timestamp":1622764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100009433","name":"Istituto Auxologico Italiano","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100009433","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Endocrinol."],"abstract":"<jats:p>The mechanisms underlying thyroid gland development have a central interest in biology and this review is aimed to provide an update on the recent advancements on the early steps of thyroid differentiation that were obtained in the zebrafish, because this teleost fish revealed to be a suitable organism to study the early developmental stages. Physiologically, the thyroid precursors fate is delineated by the appearance among the endoderm cells of the foregut of a restricted cell population expressing specific transcription factors, including <jats:italic>pax2a<\/jats:italic>, <jats:italic>nkx2.4b<\/jats:italic>, and <jats:italic>hhex<\/jats:italic>. The committed thyroid primordium first appears as a thickening of the pharyngeal floor of the anterior endoderm, that subsequently detaches from the floor and migrates to its final location where it gives rise to the thyroid hormone-producing follicles. At variance with mammalian models, thyroid precursor differentiation in zebrafish occurs early during the developmental process before the dislocation to the eutopic positioning of thyroid follicles. Several pathways have been implicated in these early events and nowadays there is evidence of a complex crosstalk between intrinsic (coming from the endoderm and thyroid precursors) and extrinsic factors (coming from surrounding tissues, as the cardiac mesoderm) whose organization in time and space is probably required for the proper thyroid development. In particular, Notch, Shh, Fgf, Bmp, and Wnt signaling seems to be required for the commitment of endodermal cells to a thyroid fate at specific developmental windows of zebrafish embryo. Here, we summarize the recent findings produced in the various zebrafish experimental models with the aim to define a comprehensive picture of such complicated puzzle.<\/jats:p>","DOI":"10.3389\/fendo.2021.664557","type":"journal-article","created":{"date-parts":[[2021,6,4]],"date-time":"2021-06-04T08:43:34Z","timestamp":1622796214000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":18,"title":["From Endoderm to Progenitors: An Update on the Early Steps of Thyroid Morphogenesis in the Zebrafish"],"prefix":"10.3389","volume":"12","author":[{"given":"Federica","family":"Marelli","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Giuditta","family":"Rurale","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Luca","family":"Persani","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2021,6,4]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.145615","article-title":"Development of the Thyroid Gland","volume":"144","author":"Nilsson","year":"2017","journal-title":"Development"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1210\/en.2011-0204","article-title":"Minireview: Intrinsic and Extrinsic Factors in Thyroid Gland Development: An Update","volume":"152","author":"De Felice","year":"2011","journal-title":"Endocrinology"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1210\/er.2003-0028","article-title":"Thyroid Development and its Disorders: Genetics and Molecular Mechanisms","volume":"25","author":"De Felice","year":"2004","journal-title":"Endocr Rev"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1242\/dev.129.15.3751","article-title":"Pax2.1 is Required for the Development of Thyroid Follicles in Zebrafish","volume":"129","author":"Wendl","year":"2002","journal-title":"Development"},{"key":"B5","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.mce.2009.12.008","article-title":"Morphogenesis of the Thyroid Gland","volume":"323","author":"Fagman","year":"2010","journal-title":"Mol Cell Endocrinol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1677\/JME-10-0084","article-title":"Morphogenetics of Early Thyroid Development","volume":"46","author":"Fagman","year":"2011","journal-title":"J Mol Endocrinol"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1093\/hmg\/ddx145","article-title":"A Frequent Oligogenic Involvement in Congenital Hypothyroidism","volume":"26","author":"de Filippis","year":"2017","journal-title":"Hum Mol Genet"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.20831","article-title":"Analysis of Origin and Growth of the Thyroid Gland in Zebrafish","volume":"235","author":"Alt","year":"2006","journal-title":"Dev Dyn"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1089\/thy.2019.0122","article-title":"Small-Molecule Screening in Zebrafish Embryos Identifies Signaling Pathways Regulating Early Thyroid Development","volume":"29","author":"Haerlingen","year":"2019","journal-title":"Thyroid"},{"key":"B10","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/j.coemr.2018.01.005","article-title":"Zebrafish Embryo: A New Model for Studying Thyroidmorphogenesis","volume":"2","author":"Larriv\u00e9e-Vanier","year":"2018","journal-title":"Curr Opin Endocrine Metab Res"},{"key":"B11","doi-asserted-by":"publisher","first-page":"2137","DOI":"10.12688\/f1000research.12142.1","article-title":"How Zebrafish Research has Helped in Understanding Thyroid Diseases","volume":"6","author":"Marelli","year":"2017","journal-title":"F1000Res"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1159\/000357079","article-title":"New Model Systems to Illuminate Thyroid Organogenesis. Part I: An Update on the Zebrafish Toolbox","volume":"2","author":"Opitz","year":"2013","journal-title":"Eur Thyroid J"},{"key":"B13","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.mce.2009.05.011","article-title":"Thyroid Gland Development and Function in the Zebrafish Model","volume":"312","author":"Porazzi","year":"2009","journal-title":"Mol Cell Endocrinol"},{"key":"B14","doi-asserted-by":"publisher","first-page":"5647","DOI":"10.1038\/s41598-018-24036-4","article-title":"A Rapid CRISPR\/Cas-Based Mutagenesis Assay in Zebrafish for Identification of Genes Involved in Thyroid Morphogenesis and Function","volume":"8","author":"Trubiroha","year":"2018","journal-title":"Sci Rep"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1089\/thy.2019.0828","article-title":"Enhanced Canonical Wnt Signaling During Early Zebrafish Development Perturbs the Interaction of Cardiac Mesoderm and Pharyngeal Endoderm and Causes Thyroid Specification Defects","volume":"31","author":"Vandernoot","year":"2021","journal-title":"Thyroid"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1101\/2020.08.13.249540","article-title":"Timed Mesodermal FGF and BMP Govern the Multistep Thyroid Specification","author":"Haerlingen","year":"2020","journal-title":"bioRxiv"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1210\/me.2011-0046","article-title":"TSH Receptor Function is Required for Normal Thyroid Differentiation in Zebrafish","volume":"25","author":"Opitz","year":"2011","journal-title":"Mol Endocrinol"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1016\/S0960-9822(00)80016-0","article-title":"A Molecular Pathway Leading to Endoderm Formation in Zebrafish","volume":"9","author":"Alexander","year":"1999","journal-title":"Curr Biol"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1242\/dev.056697","article-title":"Regulation of Endoderm Formation and Left-Right Asymmetry by miR-92 During Early Zebrafish Development","volume":"138","author":"Li","year":"2011","journal-title":"Development"},{"key":"B20","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1016\/S0925-4773(02)00327-1","article-title":"From Endoderm Formation to Liver and Pancreas Development in Zebrafish","volume":"120","author":"Ober","year":"2003","journal-title":"Mech Dev"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1016\/S0925-4773(00)00345-2","article-title":"Expression of nk2.1a During Early Development of the Thyroid Gland in Zebrafish","volume":"95","author":"Rohr","year":"2000","journal-title":"Mech Dev"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2012.09.011","article-title":"Transgenic Zebrafish Illuminate the Dynamics of Thyroid Morphogenesis and its Relationship to Cardiovascular Development","volume":"372","author":"Opitz","year":"2012","journal-title":"Dev Biol"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1242\/dev.02550","article-title":"Arteries Define the Position of the Thyroid Gland During its Developmental Relocalisation","volume":"133","author":"Alt","year":"2006","journal-title":"Development"},{"key":"B24","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/S0012-1606(03)00436-6","article-title":"Zebrafish Hhex, nk2.1a, and pax2.1 Regulate Thyroid Growth and Differentiation Downstream of Nodal-dependent Transcription Factors","volume":"263","author":"Elsalini","year":"2003","journal-title":"Dev Biol"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.4161\/cc.10.20.17764","article-title":"Endocrine Pancreas Development in Zebrafish","volume":"10","author":"Tehrani","year":"2011","journal-title":"Cell Cycle"},{"key":"B26","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1038\/nrg837","article-title":"Regulatory Phases of Early Liver Development: Paradigms of Organogenesis","volume":"3","author":"Zaret","year":"2002","journal-title":"Nat Rev Genet"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.4161\/cc.7.3.5348","article-title":"Organ Size Determination and the Limits of Regulation","volume":"7","author":"Stanger","year":"2008","journal-title":"Cell Cycle"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1038\/nature05537","article-title":"Organ Size is Limited by the Number of Embryonic Progenitor Cells in the Pancreas But Not the Liver","volume":"445","author":"Stanger","year":"2007","journal-title":"Nature"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2009.01.014","article-title":"Early Thyroid Development Requires a Tbx1-Fgf8 Pathway","volume":"328","author":"Lania","year":"2009","journal-title":"Dev Biol"},{"key":"B30","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/S1095-6433(01)00304-X","article-title":"Which Came First, the Lung or the Breath","volume":"129","author":"Perry","year":"2001","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B31","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.dci.2016.02.020","article-title":"The Zebrafish as a Model to Study Intestinal Inflammation","volume":"64","author":"Brugman","year":"2016","journal-title":"Dev Comp Immunol"},{"key":"B32","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1002\/jmor.20128","article-title":"Homology of Lungs and Gas Bladders: Insights From Arterial Vasculature","volume":"274","author":"Longo","year":"2013","journal-title":"J Morphol"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1111\/j.1440-169X.2005.00815.x","article-title":"Endoderm Development in Vertebrates: Fate Mapping, Induction and Regional Specification","volume":"47","author":"Fukuda","year":"2005","journal-title":"Dev Growth Differ"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1242\/dev.122.7.2225","article-title":"Specification of Cell Fates at the Dorsal Margin of the Zebrafish Gastrula","volume":"122","author":"Melby","year":"1996","journal-title":"Development"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1242\/dev.126.4.827","article-title":"Origin and Development of the Zebrafish Endoderm","volume":"126","author":"Warga","year":"1999","journal-title":"Development"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2008.01.028","article-title":"Live Analysis of Endodermal Layer Formation Identifies Random Walk as a Novel Gastrulation Movement","volume":"18","author":"Pezeron","year":"2008","journal-title":"Curr Biol"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/j.mod.2007.01.001","article-title":"Evolution of the Mechanisms and Molecular Control of Endoderm Formation","volume":"124","author":"Grapin-Botton","year":"2007","journal-title":"Mech Dev"},{"key":"B38","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1101\/gad.974902","article-title":"A Glimpse Into the Molecular Entrails of Endoderm Formation","volume":"16","author":"Stainier","year":"2002","journal-title":"Genes Dev"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.20905","article-title":"Retinoic Acid is Required for Endodermal Pouch Morphogenesis and Not for Pharyngeal Endoderm Specification","volume":"235","author":"Kopinke","year":"2006","journal-title":"Dev Dyn"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1242\/dev.194738","article-title":"The Second Pharyngeal Pouch is Generated by Dynamic Remodeling of Endodermal Epithelium in Zebrafish","volume":"147","author":"Okada","year":"2020","journal-title":"Development"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/bs.ctdb.2019.10.009","article-title":"Zebrafish Gastrulation: Putting Fate in Motion","volume":"136","author":"Pinheiro","year":"2020","journal-title":"Curr Top Dev Biol"},{"key":"B42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1159\/000106814","article-title":"Murine Models for the Study of Thyroid Gland Development","volume":"10","author":"Felice","year":"2007","journal-title":"Endocr Dev"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.2001.0347","article-title":"Endoderm Specification and Differentiation in Xenopus Embryos","volume":"236","author":"Horb","year":"2001","journal-title":"Dev Biol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-genet-110410-132517","article-title":"Maternal and Zygotic Control of Zebrafish Dorsoventral Axial Patterning","volume":"45","author":"Langdon","year":"2011","journal-title":"Annu Rev Genet"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/j.tcb.2010.06.009","article-title":"The Yolk Syncytial Layer in Early Zebrafish Development","volume":"20","author":"Carvalho","year":"2010","journal-title":"Trends Cell Biol"},{"key":"B46","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/bs.ctdb.2019.08.002","article-title":"Setting Up for Gastrulation in Zebrafish","volume":"136","author":"Marlow","year":"2020","journal-title":"Curr Top Dev Biol"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1242\/dev.070730","article-title":"The Hypoblast (Visceral Endoderm): An Evo-Devo Perspective","volume":"139","author":"Stern","year":"2012","journal-title":"Development"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2007.08.008","article-title":"Nodal Signals Mediate Interactions Between the Extra-Embryonic and Embryonic Tissues in Zebrafish","volume":"310","author":"Fan","year":"2007","journal-title":"Dev Biol"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1038\/26013","article-title":"Zebrafish Organizer Development and Germ-Layer Formation Require Nodal-Related Signals","volume":"395","author":"Feldman","year":"1998","journal-title":"Nature"},{"key":"B50","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1146\/annurev.genet.37.110801.143752","article-title":"Molecular Genetics of Axis Formation in Zebrafish","volume":"39","author":"Schier","year":"2005","journal-title":"Annu Rev Genet"},{"key":"B51","doi-asserted-by":"publisher","first-page":"889","DOI":"10.1152\/physrev.00038.2016","article-title":"Little Fish, Big Data: Zebrafish as a Model for Cardiovascular and Metabolic Disease","volume":"97","author":"Gut","year":"2017","journal-title":"Physiol Rev"},{"key":"B52","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1038\/35047564","article-title":"Zebrafish Genetics and Vertebrate Heart Formation","volume":"2","author":"Stainier","year":"2001","journal-title":"Nat Rev Genet"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a003459","article-title":"Nodal Morphogens","volume":"1","author":"Schier","year":"2009","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1038\/361543a0","article-title":"Nodal is a Novel TGF-beta-like Gene Expressed in the Mouse Node During Gastrulation","volume":"361","author":"Zhou","year":"1993","journal-title":"Nature"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.2001.0510","article-title":"Regulation of Nodal Signalling and Mesendoderm Formation by TARAM-A, a TGFbeta-related Type I Receptor","volume":"241","author":"Aoki","year":"2002","journal-title":"Dev Biol"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1242\/dev.02387","article-title":"Zebrafish Endoderm Formation is Regulated by Combinatorial Nodal, FGF and BMP Signalling","volume":"133","author":"Poulain","year":"2006","journal-title":"Development"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1242\/dev.128.1.125","article-title":"Multiple Roles for Gata5 in Zebrafish Endoderm Formation","volume":"128","author":"Reiter","year":"2001","journal-title":"Development"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1242\/dev.126.14.3067","article-title":"Induction of the Mesendoderm in the Zebrafish Germ Ring by Yolk Cell-Derived TGF-Beta Family Signals and Discrimination of Mesoderm and Endoderm by FGF","volume":"126","author":"Rodaway","year":"1999","journal-title":"Development"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1007\/s00427-002-0279-3","article-title":"Phenylthiourea Disrupts Thyroid Function in Developing Zebrafish","volume":"212","author":"Elsalini","year":"2003","journal-title":"Dev Genes Evol"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1242\/dev.124.2.327","article-title":"The One-Eyed Pinhead Gene Functions in Mesoderm and Endoderm Formation in Zebrafish and Interacts With No Tail","volume":"124","author":"Schier","year":"1997","journal-title":"Development"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/148.1.373","article-title":"Genetic Analysis of Chromosomal Rearrangements in the Cyclops Region of the Zebrafish Genome","volume":"148","author":"Talbot","year":"1998","journal-title":"Genetics"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1038\/350339a0","article-title":"The Cyclops Mutation Blocks Specification of the Floor Plate of the Zebrafish Central Nervous System","volume":"350","author":"Hatta","year":"1991","journal-title":"Nature"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1101\/gad.892301","article-title":"Casanova Encodes a Novel Sox-Related Protein Necessary and Sufficient for Early Endoderm Formation in Zebrafish","volume":"15","author":"Kikuchi","year":"2001","journal-title":"Genes Dev"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1101\/gad.14.10.1279","article-title":"The Zebrafish Bonnie and Clyde Gene Encodes a Mix Family Homeodomain Protein That Regulates the Generation of Endodermal Precursors","volume":"14","author":"Kikuchi","year":"2000","journal-title":"Genes Dev"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1186\/1471-213X-7-22","article-title":"Time-Dependent Patterning of the Mesoderm and Endoderm by Nodal Signals in Zebrafish","volume":"7","author":"Hagos","year":"2007","journal-title":"BMC Dev Biol"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.10483","article-title":"Notch Signaling can Regulate Endoderm Formation in Zebrafish","volume":"229","author":"Kikuchi","year":"2004","journal-title":"Dev Dyn"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1126\/science.284.5415.770","article-title":"Notch Signaling: Cell Fate Control and Signal Integration in Development","volume":"284","author":"Artavanis-Tsakonas","year":"1999","journal-title":"Science"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1210\/er.2006-0046","article-title":"Notch Signaling in Development and Cancer","volume":"28","author":"Bolos","year":"2007","journal-title":"Endocr Rev"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2009","article-title":"Notch Signalling: A Simple Pathway Becomes Complex","volume":"7","author":"Bray","year":"2006","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1677\/JOE-07-0242","article-title":"Cell and Molecular Biology of Notch","volume":"194","author":"Fiuza","year":"2007","journal-title":"J Endocrinol"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.1242\/dev.125.3.371","article-title":"Regulation of Neuronal Specification in the Zebrafish Spinal Cord by Delta Function","volume":"125","author":"Appel","year":"1998","journal-title":"Development"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1186\/1471-213X-1-13","article-title":"Delta-Notch Signaling and Lateral Inhibition in Zebrafish Spinal Cord Development","volume":"1","author":"Appel","year":"2001","journal-title":"BMC Dev Biol"},{"key":"B73","doi-asserted-by":"publisher","first-page":"17","DOI":"10.3390\/ijms17010048","article-title":"Notch Signaling in Pancreatic Development","volume":"17","author":"Li","year":"2015","journal-title":"Int J Mol Sci"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1242\/dev.013722","article-title":"Conditional Ablation of Notch Signaling in Pancreatic Development","volume":"135","author":"Nakhai","year":"2008","journal-title":"Development"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1016\/S1534-5807(03)00331-9","article-title":"Notch Signaling: A Rheostat Regulating Oligodendrocyte Differentiation","volume":"5","author":"Popko","year":"2003","journal-title":"Dev Cell"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.3389\/fnana.2014.00140","article-title":"Notch Signaling and Proneural Genes Work Together to Control the Neural Building Blocks for the Initial Scaffold in the Hypothalamus","volume":"8","author":"Ware","year":"2014","journal-title":"Front Neuroanat"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1210\/en.2011-1888","article-title":"Disruptions of Global and JAGGED1-mediated Notch Signaling Affect Thyroid Morphogenesis in the Zebrafish","volume":"153","author":"Porazzi","year":"2012","journal-title":"Endocrinology"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1210\/jc.2015-3403","article-title":"Jag1 Loss-Of-Function Variations as a Novel Predisposing Event in the Pathogenesis of Congenital Thyroid Defects","volume":"101","author":"de Filippis","year":"2016","journal-title":"J Clin Endocrinol Metab"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2004.10.027","article-title":"Differential Expression of Sonic Hedgehog Along the Anterior-Posterior Axis Regulates Patterning of Pharyngeal Pouch Endoderm and Pharyngeal Endoderm-Derived Organs","volume":"278","author":"Moore-Scott","year":"2005","journal-title":"Dev Biol"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.96.20.11376","article-title":"Multiple Left-Right Asymmetry Defects in Shh(-\/-) Mutant Mice Unveil a Convergence of the Shh and Retinoic Acid Pathways in the Control of Lefty-1","volume":"96","author":"Tsukui","year":"1999","journal-title":"Proc Natl Acad Sci USA"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.4172\/2168-9296.1000129","article-title":"Misguided Migration of C Cell Precursors to Extra-Thyroidal Locations Related to Defective Pharyngeal Pouch Development in Shh Deficient Mice","volume":"2","author":"Westerlund","year":"2013","journal-title":"Cell Dev Biol"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1101\/gad.1693608","article-title":"Hedgehog: Functions and Mechanisms","volume":"22","author":"Varjosalo","year":"2008","journal-title":"Genes Dev"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1089\/thy.2019.0196","article-title":"Glis3 as a Critical Regulator of Thyroid Primordium Specification","volume":"30","author":"Rurale","year":"2020","journal-title":"Thyroid"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2018.00730","article-title":"GLIS3 and Thyroid: A Pleiotropic Candidate Gene for Congenital Hypothyroidism","volume":"9","author":"Rurale","year":"2018","journal-title":"Front Endocrinol (Lausanne)"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1016\/j.beem.2017.04.007","article-title":"The Role of GLIS3 in Thyroid Disease as Part of a Multisystem Disorder","volume":"31","author":"Dimitri","year":"2017","journal-title":"Best Pract Res Clin Endocrinol Metab"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1038\/ng1802","article-title":"Mutations in GLIS3 are Responsible for a Rare Syndrome With Neonatal Diabetes Mellitus and Congenital Hypothyroidism","volume":"38","author":"Senee","year":"2006","journal-title":"Nat Genet"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1128\/MCB.01259-09","article-title":"Transcription Factor Glis3, a Novel Critical Player in the Regulation of Pancreatic Beta-Cell Development and Insulin Gene Expression","volume":"29","author":"Kang","year":"2009","journal-title":"Mol Cell Biol"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M111.224964","article-title":"Modulation of the Transactivation Function and Stability of Kruppel-Like Zinc Finger Protein Gli-Similar 3 (Glis3) by Suppressor of Fused","volume":"286","author":"ZeRuth","year":"2011","journal-title":"J Biol Chem"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.1016\/j.stem.2015.09.004","article-title":"Regeneration of Thyroid Function by Transplantation of Differentiated Pluripotent Stem Cells","volume":"17","author":"Kurmann","year":"2015","journal-title":"Cell Stem Cell"},{"key":"B90","doi-asserted-by":"publisher","first-page":"398","DOI":"10.1016\/j.stem.2012.01.019","article-title":"Efficient Derivation of Purified Lung and Thyroid Progenitors From Embryonic Stem Cells","volume":"10","author":"Longmire","year":"2012","journal-title":"Cell Stem Cell"},{"key":"B91","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1242\/dev.01570","article-title":"Different Thresholds of Fibroblast Growth Factors Pattern the Ventral Foregut Into Liver and Lung","volume":"132","author":"Serls","year":"2005","journal-title":"Development"},{"key":"B92","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/S0012-1606(02)00034-9","article-title":"Unique and Conserved Aspects of Gut Development in Zebrafish","volume":"255","author":"Wallace","year":"2003","journal-title":"Dev Biol"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1242\/dev.02872","article-title":"Early Developmental Specification of the Thyroid Gland Depends on Han-Expressing Surrounding Tissue and on FGF Signals","volume":"134","author":"Wendl","year":"2007","journal-title":"Development"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1242\/dev.127.12.2573","article-title":"The bHLH Transcription Factor Hand2 Plays Parallel Roles in Zebrafish Heart and Pectoral Fin Development","volume":"127","author":"Yelon","year":"2000","journal-title":"Development"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1242\/dev.129379","article-title":"Bmp Signaling Mediates Endoderm Pouch Morphogenesis by Regulating Fgf Signaling in Zebrafish","volume":"143","author":"Lovely","year":"2016","journal-title":"Development"},{"key":"B96","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.ydbio.2004.01.048","article-title":"BMP Receptor IA is Required in the Mammalian Embryo for Endodermal Morphogenesis and Ectodermal Patterning","volume":"270","author":"Davis","year":"2004","journal-title":"Dev Biol"},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2010.02.038","article-title":"Bmpr1a is Required for Proper Migration of the AVE Through Regulation of Dkk1 Expression in the Pre-Streak Mouse Embryo","volume":"341","author":"Miura","year":"2010","journal-title":"Dev Biol"},{"key":"B98","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/S0925-4773(02)00252-6","article-title":"BMP Signalling Regulates Anteroposterior Endoderm Patterning in Zebrafish","volume":"118","author":"Tiso","year":"2002","journal-title":"Mech Dev"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2007.11.004","article-title":"The BMP Signaling Gradient Patterns Dorsoventral Tissues in a Temporally Progressive Manner Along the Anteroposterior Axis","volume":"14","author":"Tucker","year":"2008","journal-title":"Dev Cell"},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.1242\/dev.150193","article-title":"Pluripotent Stem Cell Differentiation Reveals Distinct Developmental Pathways Regulating Lung- Versus Thyroid-Lineage Specification","volume":"144","author":"Serra","year":"2017","journal-title":"Development"},{"key":"B101","doi-asserted-by":"publisher","DOI":"10.1242\/dev.146589","article-title":"Wnt Signaling in Development and Tissue Homeostasis","volume":"145","author":"Steinhart","year":"2018","journal-title":"Development"},{"key":"B102","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a007955","article-title":"Wnt Signaling in Vertebrate Axis Specification","volume":"5","author":"Hikasa","year":"2013","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"B103","doi-asserted-by":"publisher","DOI":"10.15252\/embr.201845842","article-title":"New Roles for Wnt and BMP Signaling in Neural Anteroposterior Patterning","volume":"20","author":"Polevoy","year":"2019","journal-title":"EMBO Rep"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.3389\/fncel.2019.00227","article-title":"Wnt Signaling Deregulation in the Aging and Alzheimer\u2019s Brain","volume":"13","author":"Palomer","year":"2019","journal-title":"Front Cell Neurosci"},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.R600015200","article-title":"Wnt Signaling: Multiple Pathways, Multiple Receptors, and Multiple Transcription Factors","volume":"281","author":"Gordon","year":"2006","journal-title":"J Biol Chem"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1242\/dev.001230","article-title":"Repression of Wnt\/beta-catenin Signaling in the Anterior Endoderm is Essential for Liver and Pancreas Development","volume":"134","author":"McLin","year":"2007","journal-title":"Development"},{"key":"B107","doi-asserted-by":"publisher","DOI":"10.1242\/dev.055921","article-title":"Interplay Between Wnt2 and Wnt2bb Controls Multiple Steps of Early Foregut-Derived Organ Development","volume":"138","author":"Poulain","year":"2011","journal-title":"Development"}],"container-title":["Frontiers in Endocrinology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fendo.2021.664557\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,6,4]],"date-time":"2021-06-04T08:43:36Z","timestamp":1622796216000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fendo.2021.664557\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,6,4]]},"references-count":107,"alternative-id":["10.3389\/fendo.2021.664557"],"URL":"https:\/\/doi.org\/10.3389\/fendo.2021.664557","relation":{},"ISSN":["1664-2392"],"issn-type":[{"value":"1664-2392","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,4]]},"article-number":"664557"}}