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In this work, we have identified a novel MHC lineage, UEA, a complex multigene nonclassical class I family found in sharks (division Selachii) but not detected in chimaeras (subclass Holocephali) or rays (division Batoidea). This new lineage is distantly related to the previously reported nonclassical class I lineage UCA, which appears to be present only in dogfish sharks (order Squaliformes). UEA lacks conservation of the nine invariant residues in the peptide (ligand)\u2013binding regions (PBR) that bind to the N and C termini of bound peptide in most vertebrate classical class I proteins, which are replaced by relatively hydrophobic residues compared with the classical UAA. In fact, UEA and UCA proteins have the most hydrophobic-predicted PBR of all identified chondrichthyan class I molecules. UEA genes detected in the whale shark and bamboo shark genome projects are MHC linked. Consistent with UEA comprising a very large gene family, we detected weak expression in different tissues of the nurse shark via Northern blotting and RNA sequencing. UEA genes fall into three sublineages with unique characteristics in the PBR. UEA shares structural and genetic features with certain nonclassical class I genes in other vertebrates, such as the highly complex XNC nonclassical class I genes in Xenopus, and we anticipate that each shark gene, or at least each sublineage, will have a unique function, perhaps in bacterial defense.<\/jats:p>","DOI":"10.4049\/jimmunol.2000851","type":"journal-article","created":{"date-parts":[[2021,7,23]],"date-time":"2021-07-23T19:10:11Z","timestamp":1627067411000},"page":"824-836","source":"Crossref","is-referenced-by-count":13,"title":["A Highly Complex, MHC-Linked, 350 Million-Year-Old Shark Nonclassical Class I Lineage"],"prefix":"10.1093","volume":"207","author":[{"given":"Tereza","family":"Almeida","sequence":"first","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]},{"name":"Departamento de Biologia, Faculdade de Ci\u00eancias da Universidade do Porto , Porto ,","place":["Portugal"]},{"name":"Department of Microbiology and Immunology, University of Maryland Baltimore , Baltimore, MD"}]},{"given":"Yuko","family":"Ohta","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, University of Maryland Baltimore , Baltimore, MD"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2176-1488","authenticated-orcid":false,"given":"Arnaud","family":"Gaigher","sequence":"additional","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7173-2741","authenticated-orcid":false,"given":"Antonio","family":"Mu\u00f1oz-M\u00e9rida","sequence":"additional","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6593-2269","authenticated-orcid":false,"given":"Fabiana","family":"Neves","sequence":"additional","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7697-386X","authenticated-orcid":false,"given":"L Filipe C","family":"Castro","sequence":"additional","affiliation":[{"name":"Departamento de Biologia, Faculdade de Ci\u00eancias da Universidade do Porto , Porto ,","place":["Portugal"]},{"name":"Interdisciplinary Centre of Marine and Environmental Research, University of Porto , Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6857-7581","authenticated-orcid":false,"given":"Andr\u00e9 M","family":"Machado","sequence":"additional","affiliation":[{"name":"Interdisciplinary Centre of Marine and Environmental Research, University of Porto , Porto ,","place":["Portugal"]}]},{"given":"Pedro J","family":"Esteves","sequence":"additional","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]},{"name":"Departamento de Biologia, Faculdade de Ci\u00eancias da Universidade do Porto , Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3396-9822","authenticated-orcid":false,"given":"Ana","family":"Ver\u00edssimo","sequence":"additional","affiliation":[{"name":"CIBIO-InBIO, Centro de Investigac\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus Agr\u00e1rio de Vair\u00e3o, Universidade do Porto , Vair\u00e3o, Porto ,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2792-5084","authenticated-orcid":false,"given":"Martin F","family":"Flajnik","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, University of Maryland Baltimore , Baltimore, MD"}]}],"member":"286","published-online":{"date-parts":[[2021,8,1]]},"reference":[{"key":"2025022610275099600_r1","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1016\/0923-2494(96)89640-4","article-title":"Origin and evolution of the class I gene family: why are some of the mammalian class I genes encoded outside the major histocompatibility complex?","volume":"147","author":"Kasahara","year":"1996","journal-title":"Res. 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