{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T18:55:11Z","timestamp":1778180111405,"version":"3.51.4"},"reference-count":154,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2020,8,11]],"date-time":"2020-08-11T00:00:00Z","timestamp":1597104000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>The thymus generates central immune tolerance by producing self-restricted and self-tolerant T-cells as a result of interactions between the developing thymocytes and the stromal microenvironment, mainly formed by the thymic epithelial cells. The thymic epithelium derives from the endoderm of the pharyngeal pouches, embryonic structures that rely on environmental cues from the surrounding mesenchyme for its development. Here, we review the most recent advances in our understanding of the molecular mechanisms involved in early thymic organogenesis at stages preceding the expression of the transcription factor Foxn1, the early marker of thymic epithelial cells identity. Foxn1-independent developmental stages, such as the specification of the pharyngeal endoderm, patterning of the pouches, and thymus fate commitment are discussed, with a special focus on epithelial\u2013mesenchymal interactions.<\/jats:p>","DOI":"10.3390\/ijms21165765","type":"journal-article","created":{"date-parts":[[2020,8,11]],"date-time":"2020-08-11T09:28:57Z","timestamp":1597138137000},"page":"5765","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["Thymus Inception: Molecular Network in the Early Stages of Thymus Organogenesis"],"prefix":"10.3390","volume":"21","author":[{"given":"Marta","family":"Figueiredo","sequence":"first","affiliation":[{"name":"Instituto de Histologia e Biologia do Desenvolvimento, Faculdade de Medicina da Universidade de Lisboa, Edif\u00edcio Egas Moniz, Av. Prof. Egas Moniz, 1649-028 Lisboa, Portugal"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rita","family":"Zilh\u00e3o","sequence":"additional","affiliation":[{"name":"Departamento de Biologia Vegetal, Faculdade de Ci\u00eancias, Universidade de Lisboa, 1740-016 Lisboa, Portugal"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6700-4976","authenticated-orcid":false,"given":"H\u00e9lia","family":"Neves","sequence":"additional","affiliation":[{"name":"Instituto de Histologia e Biologia do Desenvolvimento, Faculdade de Medicina da Universidade de Lisboa, Edif\u00edcio Egas Moniz, Av. Prof. Egas Moniz, 1649-028 Lisboa, Portugal"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2020,8,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.immuni.2011.12.016","article-title":"Intravital Imaging of Thymopoiesis Reveals Dynamic Lympho-Epithelial Interactions","volume":"36","author":"Hess","year":"2012","journal-title":"Immunity"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.it.2012.02.005","article-title":"Evolution of lymphoid tissues","volume":"33","author":"Boehm","year":"2012","journal-title":"Trends Immunol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1017\/S0016672300010168","article-title":"\u201cNude\u201d, a new hairless gene with pleiotropic effects in the mouse","volume":"8","author":"Flanagan","year":"1966","journal-title":"Genet. Res."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"32","DOI":"10.2214\/ajr.103.1.32","article-title":"Congenital absence of the thymus","volume":"103","author":"Kirkpatrick","year":"1968","journal-title":"Am. J. Roentgenol. Radium Ther. Nucl. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1001\/archderm.141.5.647","article-title":"Nail Dystrophy Associated With a Heterozygous Mutation of the Nude\/SCID Human FOXN1 (WHN) Gene","volume":"141","author":"Auricchio","year":"2005","journal-title":"Arch. Dermatol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1016\/S0140-6736(61)90693-6","article-title":"Immunological function of the thymus","volume":"2","author":"Miller","year":"1961","journal-title":"Lance"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.1126\/science.3616623","article-title":"Tolerance Induced by Thymic Epithelial Grafts in Birds","volume":"237","author":"Ohki","year":"1987","journal-title":"Science"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"3865","DOI":"10.1242\/dev.059998","article-title":"Mechanisms of thymus organogenesis and morphogenesis","volume":"138","author":"Gordon","year":"2011","journal-title":"Development"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1007\/978-1-59745-395-0_8","article-title":"Thymus organogenesis and development of the thymic stroma","volume":"380","author":"Nowell","year":"2007","journal-title":"Methods Mol. Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1038\/nri1781","article-title":"Journey through the Thymus: Stromal Guides for T-Cell Development and Selection","volume":"6","author":"Takahama","year":"2006","journal-title":"Nat. Rev. Immunol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1016\/j.it.2009.07.010","article-title":"Thymic epithelial cells: The multi-tasking framework of the T cell \u201ccradle\u201d","volume":"30","author":"Alves","year":"2009","journal-title":"Trends Immunol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.diff.2019.06.002","article-title":"Molecular regulatory networks of thymic epithelial cell differentiation","volume":"107","author":"Luan","year":"2019","journal-title":"Differentiation"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Vaidya, H.J., Briones Leon, A., and Blackburn, C.C. (2016). FOXN1 in thymus organogenesis and development. Eur. J. Immunol., 46.","DOI":"10.1002\/eji.201545814"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Matsumoto, M., Rodrigues, P.M., Sousa, L., Tsuneyama, K., Matsumoto, M., and Alves, N.L. (2019). The Ins and Outs of Thymic Epithelial Cell Differentiation and Function. Thymus Transcriptome and Cell Biology, Springer International Publishing.","DOI":"10.1007\/978-3-030-12040-5_3"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1038\/nri.2017.12","article-title":"Generation of diversity in thymic epithelial cells","volume":"17","author":"Takahama","year":"2017","journal-title":"Nat. Rev. Immunol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"954","DOI":"10.1038\/nri2187","article-title":"Generating intrathymic microenvironments to establish T-cell tolerance","volume":"7","author":"Anderson","year":"2007","journal-title":"Nat. Rev. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2461","DOI":"10.2741\/3866","article-title":"Structure and function of the thymic microenvironment","volume":"16","author":"Manley","year":"2011","journal-title":"Front. Biosci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1093\/intimm\/4.6.707","article-title":"The thymic compartment responsible for positive selection of CD4+ T cells","volume":"4","author":"Cosgrove","year":"1992","journal-title":"Int. Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1084\/jem.20031677","article-title":"Medullary epithelial cells of the human thymus express a highly diverse selection of tissue-specific genes colocalized in chromosomal clusters","volume":"199","author":"Gotter","year":"2004","journal-title":"J. Exp. Med."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1038\/nri3667","article-title":"Positive and negative selection of the T cell repertoire: What thymocytes see and don\u2019t see","volume":"14","author":"Klein","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1111\/imr.12412","article-title":"Medullary thymic epithelial stem cells: Role in thymic epithelial cell maintenance and thymic involution","volume":"271","author":"Hamazaki","year":"2016","journal-title":"Immunol. Rev."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Irla, M. (2019). Thymic Crosstalk: An Overview of the Complex Cellular Interactions That Control the Establishment of T-Cell Tolerance. Thymus Transcriptome and Cell Biology, Springer International Publishing.","DOI":"10.1007\/978-3-030-12040-5_6"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1084\/jem.142.1.17","article-title":"Tracing of cells of the avian thymus through embryonic life in interspecific chimeras","volume":"142","author":"Jotereau","year":"1975","journal-title":"J. Exp. Med."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.ydbio.2011.10.022","article-title":"Modulation of Bmp4 signalling in the epithelial-mesenchymal interactions that take place in early thymus and parathyroid development in avian embryos","volume":"361","author":"Neves","year":"2012","journal-title":"Dev. Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/S0925-4773(01)00333-1","article-title":"Gcm2 and Foxn1 mark early parathyroid- and thymus-specific domains in the developing third pharyngeal pouch","volume":"103","author":"Gordon","year":"2001","journal-title":"Mech. Dev."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/j.semcdb.2010.01.022","article-title":"The pharyngeal pouches and clefts: Development, evolution, structure and derivatives","volume":"21","author":"Grevellec","year":"2010","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"17716","DOI":"10.1073\/pnas.0406116101","article-title":"The Origin of the Parathyroid Gland","volume":"101","author":"Okabe","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_28","unstructured":"Ashford, M. (2014). Development of Parathyroid Glands and C-Cells. Parathyroid Glands: Regulation, Role in Human Disease and Indications for Surgery, Nova Science."},{"key":"ref_29","first-page":"589","article-title":"3D immunofluorescence analysis of early thymic morphogenesis and medulla development","volume":"30","author":"Cejalvo","year":"2015","journal-title":"Histol. Histopathol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/0012-1606(60)90009-9","article-title":"Morphogenetic interactions in the development of the mouse thymus gland","volume":"2","author":"Auerbach","year":"1960","journal-title":"Dev. Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"498","DOI":"10.1126\/science.6606851","article-title":"Dependence of Thymus Development on Derivatives of the Neural Crest","volume":"223","author":"Bockman","year":"1984","journal-title":"Science"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1111\/j.1469-7580.2005.00472.x","article-title":"The role of the endoderm in the development and evolution of the pharyngeal arches","volume":"207","author":"Graham","year":"2005","journal-title":"J. Anat."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"546","DOI":"10.1038\/ni1064","article-title":"Functional evidence for a single endodermal origin for the thymic epithelium","volume":"5","author":"Gordon","year":"2004","journal-title":"Nat. Immunol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1016\/j.ydbio.2008.12.019","article-title":"Increased thymus- and decreased parathyroid-fated organ domains in Splotch mutant embryos","volume":"327","author":"Griffith","year":"2009","journal-title":"Dev Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"10555","DOI":"10.1073\/pnas.1005129107","article-title":"Mouse and zebrafish Hoxa3 Orthologues Have Nonequivalent In Vivo Protein Function","volume":"107","author":"Chen","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Chojnowski, J.L., Masuda, K., Trau, H.A., Thomas, K., Capecchi, M., and Manley, N.R. (2014). Multiple roles for HOXA3 in regulating thymus and parathyroid differentiation and morphogenesis in mouse. Development, 141.","DOI":"10.1242\/dev.110833"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1002\/aja.1001950404","article-title":"A series of normal stages in the development of the chick embryo. 1951","volume":"195","author":"Hamburger","year":"1992","journal-title":"Dev. Dyn."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"5344","DOI":"10.4049\/jimmunol.180.8.5344","article-title":"Neural Crest Origin of Perivascular Mesenchyme in the Adult Thymus","volume":"180","author":"Muller","year":"2008","journal-title":"J. Immunol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3183","DOI":"10.4049\/jimmunol.180.5.3183","article-title":"Contribution of Neural Crest-Derived Cells in the Embryonic and Adult Thymus","volume":"180","author":"Foster","year":"2008","journal-title":"J. Immunol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.ydbio.2016.05.010","article-title":"Temporal and spatial requirements for Hoxa3 in mouse embryonic development","volume":"415","author":"Chojnowski","year":"2016","journal-title":"Dev. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"13414","DOI":"10.1073\/pnas.1003747107","article-title":"EphB-Ephrin-B2 Interactions are Required for Thymus Migration during Organogenesis","volume":"107","author":"Foster","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1016\/0167-5699(94)90246-1","article-title":"Crosstalk in the mouse thymus","volume":"15","author":"Shores","year":"1994","journal-title":"Trends Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"11822","DOI":"10.1073\/pnas.95.20.11822","article-title":"Interdependence of Cortical Thymic Epithelial Cell Differentiation and T-Lineage Commitment","volume":"95","author":"Klug","year":"1998","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1583","DOI":"10.1242\/dev.127.8.1583","article-title":"Stepwise development of thymic microenvironments in vivo is regulated by thymocyte subsets","volume":"127","author":"Terhorst","year":"2000","journal-title":"Development"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1111\/j.0105-2896.2006.00347.x","article-title":"Establishment and functioning of intrathymic microenvironments","volume":"209","author":"Anderson","year":"2006","journal-title":"Immunol. Rev."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1146\/annurev-immunol-051116-052320","article-title":"Thymic Epithelial Cells","volume":"35","author":"Abramson","year":"2017","journal-title":"Annu. Rev. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1002\/eji.201747006","article-title":"Dissecting and modeling the emergent murine TEC compartment during ontogeny","volume":"47","author":"Brunk","year":"2017","journal-title":"Eur. J. Immunol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/j.ydbio.2007.02.014","article-title":"Gcm2 is required for the differentiation and survival of parathyroid precursor cells in the parathyroid\/thymus primordia","volume":"305","author":"Liu","year":"2007","journal-title":"Dev. Biol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1126\/science.272.5263.886","article-title":"Two genetically separable steps in the differentiation of thymic epithelium","volume":"272","author":"Nehls","year":"1996","journal-title":"Science"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Mu\u00f1oz, J.J., and Zapata, A.G. (2019). Thymus Ontogeny and Development. Thymus Transcriptome and Cell Biology, Springer International Publishing.","DOI":"10.1007\/978-3-030-12040-5_2"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1038\/372103a0","article-title":"New member of the winged-helix protein family disrupted in mouse and rat nude mutations","volume":"372","author":"Nehls","year":"1994","journal-title":"Nature"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1203","DOI":"10.1093\/intimm\/13.9.1203","article-title":"Two distinct steps of immigration of hematopoietic progenitors into the early thymus anlage","volume":"13","author":"Itoi","year":"2001","journal-title":"Int. Immunol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"992","DOI":"10.1038\/nature04850","article-title":"Formation of a functional thymus initiated by a postnatal epithelial progenitor cell","volume":"441","author":"Bleul","year":"2006","journal-title":"Nature"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1106","DOI":"10.1016\/j.molimm.2009.10.034","article-title":"Thymic cysts originate from Foxn1 positive thymic medullary epithelium","volume":"47","author":"Vroegindeweij","year":"2010","journal-title":"Mol. Immunol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1007\/s00418-019-01818-z","article-title":"FoxN1 mediates thymic cortex\u2013medulla differentiation through modifying a developmental pattern based on epithelial tubulogenesis","volume":"152","author":"Tobajas","year":"2019","journal-title":"Histochem. Cell Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"14912","DOI":"10.1073\/pnas.93.25.14912","article-title":"The Gcm-Motif: A Novel DNA-Binding Motif Conserved in Drosophila and Mammals","volume":"93","author":"Akiyama","year":"1996","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1038\/35018111","article-title":"Genetic ablation of parathyroid glands reveals another source of parathyroid hormone","volume":"406","author":"Chen","year":"2000","journal-title":"Nature"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/S0925-4773(02)00345-3","article-title":"Aortic arch and pharyngeal phenotype in the absence of BMP-dependent neural crest in the mouse","volume":"119","author":"Papaioannou","year":"2002","journal-title":"Mech. Dev."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"5213","DOI":"10.4049\/jimmunol.175.8.5213","article-title":"BMP signaling is required for normal thymus development","volume":"175","author":"Bleul","year":"2005","journal-title":"J. Immunol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1016\/j.modgep.2006.01.011","article-title":"Bmp4 and Noggin Expression during Early Thymus and Parathyroid Organogenesis","volume":"6","author":"Patel","year":"2006","journal-title":"Gene Expr. Patterns GEP"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"5272","DOI":"10.4049\/jimmunol.181.8.5272","article-title":"Maintenance of Thymic Epithelial Phenotype Requires Extrinsic Signals in Mouse and Zebrafish","volume":"181","author":"Bleul","year":"2008","journal-title":"J. Immunol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/j.ydbio.2009.12.026","article-title":"Evidence for an early role for BMP4 signaling in thymus and parathyroid morphogenesis","volume":"339","author":"Gordon","year":"2010","journal-title":"Dev. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"8492","DOI":"10.1038\/s41598-017-09213-1","article-title":"Cooperative interaction of BMP signalling and Foxn1 gene dosage determines the size of the functionally active thymic epithelial compartment","volume":"7","author":"Swann","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"3033","DOI":"10.1242\/dev.129.13.3033","article-title":"Eya1 is required for the morphogenesis of mammalian thymus, parathyroid and thyroid","volume":"129","author":"Xu","year":"2002","journal-title":"Development"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1016\/j.ydbio.2005.12.015","article-title":"Patterning of the third pharyngeal pouch into thymus\/parathyroid by Six and Eya1","volume":"293","author":"Zou","year":"2006","journal-title":"Dev. Biol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"4613","DOI":"10.1242\/dev.129.19.4613","article-title":"Fgf8 is required for pharyngeal arch and cardiovascular development in the mouse","volume":"129","author":"Smyth","year":"2002","journal-title":"Development"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"4591","DOI":"10.1242\/dev.129.19.4591","article-title":"An Fgf8 mouse mutant phenocopies human 22q11 deletion syndrome","volume":"129","author":"Frank","year":"2002","journal-title":"Development"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"6361","DOI":"10.1242\/dev.00850","article-title":"Ablation of Specific Expression Domains Reveals Discrete Functions of Ectoderm-And Endoderm-Derived FGF8 during Cardiovascular and Pharyngeal Development","volume":"130","author":"Macatee","year":"2003","journal-title":"Development"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"3456","DOI":"10.1242\/dev.079400","article-title":"Localised inhibition of FGF signalling in the third pharyngeal pouch is required for normal thymus and parathyroid organogenesis","volume":"139","author":"Gardiner","year":"2012","journal-title":"Development"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Hasten, E., and Morrow, B.E. (2019). Tbx1 and Foxi3 genetically interact in the pharyngeal pouch endoderm in a mouse model for 22q11.2 deletion syndrome. PLoS Genet., 15.","DOI":"10.1371\/journal.pgen.1008301"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"5742","DOI":"10.1073\/pnas.93.12.5742","article-title":"The Nu Gene Acts Cell-Autonomously and Is Required for Differentiation of Thymic Epithelial Progenitors","volume":"93","author":"Blackburn","year":"1996","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1128","DOI":"10.1038\/ni983","article-title":"A domain of Foxn1 required for crosstalk-dependent thymic epithelial cell differentiation","volume":"4","author":"Su","year":"2003","journal-title":"Nat. Immunol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"2144","DOI":"10.1172\/JCI42021","article-title":"Gata3-deficient mice develop parathyroid abnormalities due to dysregulation of the parathyroid-specific transcription factor Gcm2","volume":"120","author":"Grigorieva","year":"2010","journal-title":"J. Clin. Investig."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Wei, Q., and Condie, B.G. (2011). A focused In Situ Hybridization Screen Identifies Candidate Transcriptional Regulators of Thymic Epithelial Cell Development and Function. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0026795"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1016\/j.ydbio.2016.08.012","article-title":"Notch and Hedgehog in the thymus\/parathyroid common primordium: Crosstalk in organ formation","volume":"418","author":"Figueiredo","year":"2016","journal-title":"Dev. Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1989","DOI":"10.1242\/dev.121.7.1989","article-title":"The role of Hoxa-3 in mouse thymus and thyroid development","volume":"121","author":"Manley","year":"1995","journal-title":"Development"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1006\/dbio.1997.8827","article-title":"Hox group 3 paralogs regulate the development and migration of the thymus, thyroid, and parathyroid glands","volume":"195","author":"Manley","year":"1998","journal-title":"Dev. Biol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1006\/dbio.2001.0342","article-title":"Hoxa3 and pax1 regulate epithelial cell death and proliferation during thymus and parathyroid organogenesis","volume":"236","author":"Su","year":"2001","journal-title":"Dev. Biol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1242\/dev.122.1.23","article-title":"Pax1 is expressed during development of the thymus epithelium and is required for normal T-cell maturation","volume":"122","author":"Wallin","year":"1996","journal-title":"Development"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"2735","DOI":"10.1101\/gad.12.17.2735","article-title":"Pax9-deficient mice lack pharyngeal pouch derivatives and teeth and exhibit craniofacial and limb abnormalities","volume":"12","author":"Peters","year":"1998","journal-title":"Genes Dev."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1175","DOI":"10.1002\/1521-4141(200204)32:4<1175::AID-IMMU1175>3.0.CO;2-U","article-title":"Thymopoiesis requires Pax9 function in thymic epithelial cells","volume":"32","author":"Schorpp","year":"2002","journal-title":"Eur. J. Immunol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1553","DOI":"10.1242\/dev.127.8.1553","article-title":"Retinoid signaling is essential for patterning the endoderm of the third and fourth pharyngeal arches","volume":"127","author":"Wendling","year":"2000","journal-title":"Development"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1002\/dvdy.10137","article-title":"Deficits in the posterior pharyngeal endoderm in the absence of retinoids","volume":"225","author":"Quinlan","year":"2002","journal-title":"Dev. Dyn."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"2525","DOI":"10.1242\/dev.00463","article-title":"The regional pattern of retinoic acid synthesis by RALDH2 is essential for the development of posterior pharyngeal arches and the enteric nervous system","volume":"130","author":"Niederreither","year":"2003","journal-title":"Development"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1002\/dvdy.10292","article-title":"Retinoic acid signalling centres in the avian embryo identified by sites of expression of synthesising and catabolising enzymes","volume":"227","author":"Blentic","year":"2003","journal-title":"Dev. Dyn."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"2296","DOI":"10.4049\/jimmunol.172.4.2296","article-title":"Reduced Thymocyte Development in Sonic Hedgehog Knockout Embryos","volume":"172","author":"Shah","year":"2004","journal-title":"J. Immunol. (Baltim. MD 1950)"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"323","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":"Manley","year":"2005","journal-title":"Dev. Biol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1016\/j.ydbio.2011.02.012","article-title":"Shh signalling restricts the expression of Gcm2 and controls the position of the developing parathyroids","volume":"353","author":"Grevellec","year":"2011","journal-title":"Dev. Biol."},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Bain, V.E., Gordon, J., O\u2019Neil, J.D., Ramos, I., Richie, E.R., and Manley, N.R. (2016). Tissue-specific roles for sonic hedgehog signaling in establishing thymus and parathyroid organ fate. Development, 143.","DOI":"10.1242\/dev.141903"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1038\/85845","article-title":"DiGeorge syndrome phenotype in mice mutant for the T-box gene, Tbx1","volume":"27","author":"Jerome","year":"2001","journal-title":"Nat. Genet."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"928","DOI":"10.1002\/dvdy.20268","article-title":"Retinoic acid down-regulates Tbx1 expression in vivo and in vitro","volume":"232","author":"Roberts","year":"2005","journal-title":"Dev. Dyn."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1102","DOI":"10.1038\/ni850","article-title":"Wnt glycoproteins regulate the expression of FoxN1, the gene defective in nude mice","volume":"3","author":"Balciunaite","year":"2002","journal-title":"Nat. Immunol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"2695","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":"ref_94","doi-asserted-by":"crossref","unstructured":"Diman, N.Y.S.G., Remacle, S., Bertrand, N., Picard, J.J., Zaffran, S., and Rezsohazy, R. (2011). A retinoic acid responsive Hoxa3 transgene expressed in embryonic pharyngeal endoderm, cardiac neural crest and a subdomain of the second heart field. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0027624"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"5051","DOI":"10.1242\/dev.126.22.5051","article-title":"Key roles of retinoic acid receptors alpha and beta in the patterning of the caudal hindbrain, pharyngeal arches and otocyst in the mouse","volume":"126","author":"Ghyselinck","year":"1999","journal-title":"Development"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1146\/annurev.genet.39.073003.105925","article-title":"T-Box Genes in Vertebrate Development","volume":"39","author":"Naiche","year":"2005","journal-title":"Annu. Rev. Genet."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1038\/35065105","article-title":"Tbx1 haploinsufficieny in the DiGeorge syndrome region causes aortic arch defects in mice","volume":"410","author":"Lindsay","year":"2001","journal-title":"Nature"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/j.gde.2005.03.001","article-title":"Dissecting contiguous gene defects: TBX1","volume":"15","author":"Baldini","year":"2005","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"915","DOI":"10.1093\/hmg\/11.8.915","article-title":"Tbx1 mutation causes multiple cardiovascular defects and disrupts neural crest and cranial nerve migratory pathways","volume":"11","author":"Vitelli","year":"2002","journal-title":"Hum. Mol. Genet."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1006\/dbio.2001.0283","article-title":"Tbx1, a DiGeorge syndrome candidate gene, is regulated by sonic hedgehog during pharyngeal arch development","volume":"235","author":"Garg","year":"2001","journal-title":"Dev. Biol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"5307","DOI":"10.1242\/dev.02086","article-title":"Tbx1 expression in pharyngeal epithelia is necessary for pharyngeal arch artery development","volume":"132","author":"Zhang","year":"2005","journal-title":"Development"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"3583","DOI":"10.1242\/dev.111740","article-title":"Tbx1 controls the morphogenesis of pharyngeal pouch epithelia through mesodermal Wnt11r and Fgf8a","volume":"141","author":"Choe","year":"2014","journal-title":"Development"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"4387","DOI":"10.1242\/dev.02018","article-title":"Timed mutation and cell-fate mapping reveal reiterated roles of Tbx1 during embryogenesis, and a crucial function during segmentation of the pharyngeal system via regulation of endoderm expansion","volume":"132","author":"Xu","year":"2005","journal-title":"Development"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1242\/dev.02264","article-title":"Inactivation of Tbx1 in the pharyngeal endoderm results in 22q11DS malformations","volume":"133","author":"Arnold","year":"2006","journal-title":"Development"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1002\/dvdy.24147","article-title":"Endoderm-specific deletion of Tbx1 reveals an FGF-independent role for Tbx1 in pharyngeal apparatus morphogenesis","volume":"243","author":"Jackson","year":"2014","journal-title":"Dev. Dyn."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"3587","DOI":"10.1242\/dev.02539","article-title":"Mesodermal expression of Tbx1 is necessary and sufficient for pharyngeal arch and cardiac outflow tract development","volume":"133","author":"Zhang","year":"2006","journal-title":"Development"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"2950","DOI":"10.1242\/dev.111641","article-title":"Ectopic TBX1 suppresses thymic epithelial cell differentiation and proliferation during thymus organogenesis","volume":"141","author":"Reeh","year":"2014","journal-title":"Development"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"3731","DOI":"10.1242\/dev.037689","article-title":"FGF signalling: Diverse roles during early vertebrate embryogenesis","volume":"137","author":"Dorey","year":"2010","journal-title":"Development"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1242\/dev.121.2.439","article-title":"The mouse Fgf8 gene encodes a family of polypeptides and is expressed in regions that direct outgrowth and patterning in the developing embryo","volume":"121","author":"Crossley","year":"1995","journal-title":"Development"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1038\/ng0298-136","article-title":"An Fgf8 mutant allelic series generated by Cre- and Flp-mediated recombination","volume":"18","author":"Meyers","year":"1998","journal-title":"Nat. Genet."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"2419","DOI":"10.1242\/dev.02367","article-title":"Required, tissue-specific roles for Fgf8 in outflow tract formation and remodeling","volume":"133","author":"Park","year":"2006","journal-title":"Development"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1387\/ijdb.180284jg","article-title":"Hox genes in the pharyngeal region: How Hoxa3 controls early embryonic development of the pharyngeal organs","volume":"62","author":"Gordon","year":"2018","journal-title":"Int. J. Dev. Biol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"60","DOI":"10.4049\/jimmunol.174.1.60","article-title":"Ontogeny and regulation of IL-7-expressing thymic epithelial cells","volume":"174","author":"Zamisch","year":"2005","journal-title":"J. Immunol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/0092-8674(94)90290-9","article-title":"Hox genes in vertebrate development","volume":"78","author":"Krumlauf","year":"1994","journal-title":"Cell"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1146\/annurev.cellbio.042308.113423","article-title":"Hox Genes and Segmentation of the Hindbrain and Axial Skeleton","volume":"25","author":"Alexander","year":"2009","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/S1877-1173(10)92005-X","article-title":"Transcriptional regulation of thymus organogenesis and thymic epithelial cell differentiation","volume":"Volume 92","author":"Manley","year":"2010","journal-title":"Progress in Molecular Biology and Translational Science"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1016\/0092-8674(93)90115-7","article-title":"The eyes absent gene: Genetic control of cell survival and differentiation in the developing Drosophila eye","volume":"72","author":"Bonini","year":"1993","journal-title":"Cell"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1038\/nature02083","article-title":"Eya protein phosphatase activity regulates Six1-Dach-Eya transcriptional effects in mammalian organogenesis","volume":"426","author":"Li","year":"2003","journal-title":"Nature"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"1137","DOI":"10.1093\/genetics\/138.4.1137","article-title":"Sine oculis is a homeobox gene required for Drosophila visual system development","volume":"138","author":"Serikaku","year":"1994","journal-title":"Genetics"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1002\/bies.950190905","article-title":"Pax genes and organogenesis","volume":"19","author":"Dahl","year":"1997","journal-title":"Bioessays"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1006\/dbio.1995.1248","article-title":"Characterization and developmental expression of Pax9, a paired-box-containing gene related to Pax1","volume":"170","author":"Koseki","year":"1995","journal-title":"Dev. Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"1426","DOI":"10.1210\/jc.2007-1783","article-title":"Analysis of the GCM2 gene in isolated hypoparathyroidism: A molecular and biochemical study","volume":"93","author":"Maret","year":"2008","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"5399","DOI":"10.1242\/dev.126.23.5399","article-title":"Pax1 and Pax9 synergistically regulate vertebral column development","volume":"126","author":"Peters","year":"1999","journal-title":"Development"},{"key":"ref_124","unstructured":"Kelly, M. (2012). Molecular Regulation of Thymic Epithelial Lineage Specification. [Ph.D. Thesis, University of Edinburgh]."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1002\/dvdy.20160","article-title":"Expression of mouse Foxi class genes in early craniofacial development","volume":"231","author":"Ohyama","year":"2004","journal-title":"Dev. Dyn."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1084\/jem.181.4.1519","article-title":"Lymphopenia in interleukin (IL)-7 gene-deleted mice identifies IL-7 as a nonredundant cytokine","volume":"181","author":"Vieira","year":"1995","journal-title":"J. Exp. Med."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"2874","DOI":"10.1128\/MCB.20.8.2874-2879.2000","article-title":"Phenotypic characterization of the murine Nkx2.6 homeobox gene by gene targeting","volume":"20","author":"Tanaka","year":"2000","journal-title":"Mol. Cell. Biol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1084\/jem.20022135","article-title":"Differential requirement for mesenchyme in the proliferation and maturation of thymic epithelial progenitors","volume":"198","author":"Jenkinson","year":"2003","journal-title":"J. Exp. Med."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"3459","DOI":"10.1002\/dvdy.21364","article-title":"FGFR2IIIb signaling regulates thymic epithelial differentiation","volume":"236","author":"Dooley","year":"2007","journal-title":"Dev. Dyn."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"1954","DOI":"10.4049\/jimmunol.167.4.1954","article-title":"Development of the thymus requires signaling through the fibroblast growth factor receptor R2-IIIb","volume":"167","author":"Revest","year":"2001","journal-title":"J. Immunol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/s00441-013-1649-z","article-title":"Hes1 is required for the development of pharyngeal organs and survival of neural crest-derived mesenchymal cells in pharyngeal arches","volume":"353","author":"Kameda","year":"2013","journal-title":"Cell Tissue Res."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.gendis.2014.07.005","article-title":"Bone Morphogenetic Protein (BMP) signaling in development and human diseases","volume":"1","author":"Wang","year":"2014","journal-title":"Genes Dis."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"3947","DOI":"10.1182\/blood-2003-05-1657","article-title":"BMP4 acts upstream of FGF in modulating thymic stroma and regulating thymopoiesis","volume":"102","author":"Tsai","year":"2003","journal-title":"Blood"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1101\/gad.1886910","article-title":"Control of hair follicle cell fate by underlying mesenchyme through a CSL-Wnt5a-FoxN1 regulatory axis","volume":"24","author":"Hu","year":"2010","journal-title":"Genes Dev."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"6664","DOI":"10.1093\/emboj\/19.24.6664","article-title":"Inhibition of Bmp signaling affects growth and differentiation in the anagen hair follicle","volume":"19","author":"Kulessa","year":"2000","journal-title":"EMBO J."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1242\/dev.01074","article-title":"Notch signaling: Control of cell communication and cell fate","volume":"131","author":"Lai","year":"2004","journal-title":"Development"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"678","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":"ref_138","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1016\/j.ceb.2007.02.012","article-title":"Crossing paths with Notch in the hyper-network","volume":"19","author":"Hurlbut","year":"2007","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_139","first-page":"2135","article-title":"Notch signaling at a glance","volume":"126","author":"Hori","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1002\/dvdy.24291","article-title":"Otic placode cell specification and proliferation are regulated by Notch signaling in avian development","volume":"244","author":"Shida","year":"2015","journal-title":"Dev. Dyn."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"991","DOI":"10.1084\/jem.194.7.991","article-title":"Differential effects of Notch ligands Delta-1 and Jagged-1 in human lymphoid differentiation","volume":"194","author":"Jaleco","year":"2001","journal-title":"J. Exp. Med."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1146\/annurev.immunol.23.021704.115747","article-title":"Regulation of lymphoid development, differentiation, and function by the Notch pathway","volume":"23","author":"Maillard","year":"2005","journal-title":"Annu. Rev. Immunol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.immuni.2010.01.004","article-title":"Notch signaling in the immune system","volume":"32","author":"Radtke","year":"2010","journal-title":"Immunity"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"4017","DOI":"10.4049\/jimmunol.1302259","article-title":"An Overview of the Intrathymic Intricacies of T Cell Development","volume":"192","author":"Shah","year":"2014","journal-title":"J. Immunol."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/j.ydbio.2010.01.020","article-title":"Hes1 expression is reduced in Tbx1 null cells and is required for the development of structures affected in 22q11 deletion syndrome","volume":"340","author":"Papangeli","year":"2010","journal-title":"Dev. Biol."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1038\/35019553","article-title":"Fringe differentially modulates Jagged1 and Delta1 signalling through Notch1 and Notch2","volume":"2","author":"Hicks","year":"2000","journal-title":"Nat. Cell Biol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1242\/dev.120154","article-title":"Hedgehog signalling","volume":"143","author":"Lee","year":"2016","journal-title":"Development"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"899","DOI":"10.1242\/dev.006742","article-title":"Oscillatory lunatic fringe activity is crucial for segmentation of the anterior but not posterior skeleton","volume":"135","author":"Shifley","year":"2008","journal-title":"Development"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.ydbio.2006.11.009","article-title":"Sonic hedgehog in the pharyngeal endoderm controls arch pattern via regulation of Fgf8 in head ectoderm","volume":"303","author":"Haworth","year":"2007","journal-title":"Dev. Biol."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1101\/gad.1048903","article-title":"Tbx1 is regulated by tissue-specific forkhead proteins through a common Sonic hedgehog-responsive enhancer","volume":"17","author":"Yamagishi","year":"2003","journal-title":"Genes Dev."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1016\/j.ydbio.2004.08.030","article-title":"Abnormalities of caudal pharyngeal pouch development in Pbx1 knockout mice mimic loss of Hox3 paralogs","volume":"276","author":"Manley","year":"2004","journal-title":"Dev. Biol."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.3389\/fimmu.2019.01473","article-title":"Emerging roles for Eph receptors and ephrin ligands in immunity","volume":"10","author":"Darling","year":"2019","journal-title":"Front. Immunol."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1159\/000329490","article-title":"Eph\/ephrin-mediated interactions in the thymus","volume":"18","author":"Cejalvo","year":"2011","journal-title":"Neuroimmunomodulation"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1007\/s00418-014-1296-9","article-title":"Conditioned deletion of ephrinB1 and\/or ephrinB2 in either thymocytes or thymic epithelial cells alters the organization of thymic medulla and favors the appearance of thymic epithelial cysts","volume":"143","author":"Cejalvo","year":"2015","journal-title":"Histochem. Cell Biol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/21\/16\/5765\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T09:59:17Z","timestamp":1760176757000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/21\/16\/5765"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,8,11]]},"references-count":154,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2020,8]]}},"alternative-id":["ijms21165765"],"URL":"https:\/\/doi.org\/10.3390\/ijms21165765","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,8,11]]}}}