{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:05:44Z","timestamp":1792782344206,"version":"4.5.0"},"reference-count":106,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2019,1,24]],"date-time":"2019-01-24T00:00:00Z","timestamp":1548288000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"},{"start":{"date-parts":[[2019,1,24]],"date-time":"2019-01-24T00:00:00Z","timestamp":1548288000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2019,1,24]],"date-time":"2019-01-24T00:00:00Z","timestamp":1548288000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"name":"CONACyT","award":["#166024"],"award-info":[{"award-number":["#166024"]}]},{"name":"CONACyT","award":["#253274"],"award-info":[{"award-number":["#253274"]}]},{"name":"CONACyT","award":["#166024"],"award-info":[{"award-number":["#166024"]}]},{"name":"CONACyT","award":["#253274"],"award-info":[{"award-number":["#253274"]}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2019,4,19]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Since its discovery, over 30 years ago, CD5 has been used as a marker to identify T cells, B1-a cells, and B cell chronic lymphocytic leukemia cells. Throughout the years, many studies have described the functional relevance of CD5 as a modulator of T and B cell receptor signaling. However, it has not been until recent years that CD5 has emerged as a functional receptor in other areas of the immune system. Here, we review some of the most important aspects of CD5 as a modulator of TCR and BCR signaling, cell survival receptor both in T and B cells during health and disease, as well as the newly discovered roles of this receptor in thymocyte selection, T cell effector differentiation, and immune tolerance. CD5 was found to promote T cell survival by protecting autoreactive T cell from activation-induced cell death, to promote de novo induction of regulatory T cells in the periphery, to modulate Th17 and Th2 differentiation, and to modulate immune responses by modulating dendritic cell functions. CD5 is overexpressed in Tregs and Bregs, which are fundamental to maintain immune homeostasis. The newly established roles of CD5 in modulating different aspects of immune responses identify this receptor as an immune checkpoint modulator, and therefore it could be used as a target for immune intervention in different pathologies such as cancer, autoimmune diseases or infections.<\/jats:p>","DOI":"10.1002\/jlb.mr0618-226r","type":"journal-article","created":{"date-parts":[[2019,1,24]],"date-time":"2019-01-24T17:52:20Z","timestamp":1548352340000},"page":"891-904","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":92,"title":["The multiple faces of CD5"],"prefix":"10.1093","volume":"105","author":[{"given":"Erica","family":"Burgue\u00f1o-Bucio","sequence":"first","affiliation":[{"name":"Department of Immunology, Instituto de Investigaciones Biom\u00e9dicas, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Mexico City, Mexico"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carlos A","family":"Mier-Aguilar","sequence":"additional","affiliation":[{"name":"Department of Immunology, Instituto de Investigaciones Biom\u00e9dicas, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Mexico City, Mexico"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gloria","family":"Soldevila","sequence":"additional","affiliation":[{"name":"Department of Immunology, Instituto de Investigaciones Biom\u00e9dicas, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Mexico City, Mexico"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2019,1,24]]},"reference":[{"key":"2023032406175973400_jlb10304-cit-0001","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.4049\/jimmunol.123.3.1312","article-title":"A monoclonal antibody with selective reactivity with functionally mature human thymocytes and all peripheral human T cells?","volume":"123","author":"Reinherz","year":"1979","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0002","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1084\/jem.152.1.229","article-title":"An antigen shared by a human T cell subset and B cell chronic lymphocytic leukemic cells. Distribution on normal and malignant lymphoid cells?","volume":"152","author":"Boumsell","year":"1980","journal-title":"J Exp Med"},{"key":"2023032406175973400_jlb10304-cit-0003","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1016\/j.coi.2011.03.003","article-title":"The immunomodulatory properties of the CD5 lymphocyte receptor in health and disease?","volume":"23","author":"Soldevila","year":"2011","journal-title":"Curr Opin Immunol"},{"key":"2023032406175973400_jlb10304-cit-0004","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1084\/jem.152.2.280","article-title":"T cell subsets defined by expression of Lyt-1,2,3 and Thy-1 antigens. Two-parameter immunofluorescence and cytotoxicity analysis with monoclonal antibodies modifies current views?","volume":"152","author":"Ledbetter","year":"1980","journal-title":"J Exp Med"},{"key":"2023032406175973400_jlb10304-cit-0005","doi-asserted-by":"crossref","first-page":"1531","DOI":"10.4049\/jimmunol.134.3.1531","article-title":"Human B cell development. II. Subpopulations in the human fetus?","volume":"134","author":"Bofill","year":"1985","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0006","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1146\/annurev.immunol.20.100301.064833","article-title":"Origins and functions of B-1 cells with notes on the role of CD5?","volume":"20","author":"Berland","year":"2002","journal-title":"Annu Rev Immunol"},{"key":"2023032406175973400_jlb10304-cit-0007","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1111\/j.0105-2896.2004.0109.x","article-title":"Peripheral development of B cells in mouse and man?","volume":"197","author":"Carsetti","year":"2004","journal-title":"Immunol Rev"},{"key":"2023032406175973400_jlb10304-cit-0008","doi-asserted-by":"crossref","first-page":"3858","DOI":"10.4049\/jimmunol.149.12.3858","article-title":"CD5+ B cells predominate in peripheral tissues of rabbit?","volume":"149","author":"Raman","year":"1992","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0009","doi-asserted-by":"crossref","first-page":"1553","DOI":"10.4049\/jimmunol.1600193","article-title":"Human blood CD1c+ dendritic cells encompass CD5high and CD5low subsets that differ significantly in phenotype, gene expression, and functions?","volume":"198","author":"Yin","year":"2017","journal-title":"J Immunol"},{"issue":"18","key":"2023032406175973400_jlb10304-cit-0010","doi-asserted-by":"crossref","DOI":"10.1172\/jci.insight.96101","article-title":"A type of human skin dendritic cell marked by CD5 is associated with the development of inflammatory skin disease?","volume":"2","author":"Korenfeld","year":"2017","journal-title":"JCI Insight"},{"key":"2023032406175973400_jlb10304-cit-0011","doi-asserted-by":"crossref","first-page":"1988","DOI":"10.1073\/pnas.1610630114","article-title":"A distinct subset of plasmacytoid dendritic cells induces activation and differentiation of B and T lymphocytes?","volume":"114","author":"Zhang","year":"2017","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023032406175973400_jlb10304-cit-0012","doi-asserted-by":"crossref","first-page":"2301","DOI":"10.1084\/jem.188.12.2301","article-title":"CD5 expression is developmentally regulated by T cell receptor (TCR) signals and TCR avidity?","volume":"188","author":"Azzam","year":"1998","journal-title":"J Exp Med"},{"key":"2023032406175973400_jlb10304-cit-0013","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1038\/323346a0","article-title":"Isolation of complementary DNA clones encoding the human lymphocyte glycoprotein T1\/Leu-1?","volume":"323","author":"Jones","year":"1986","journal-title":"Nature"},{"key":"2023032406175973400_jlb10304-cit-0014","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1073\/pnas.84.1.204","article-title":"Molecular cloning of Ly-1, a membrane glycoprotein of mouse T lymphocytes and a subset of B cells: molecular homology to its human counterpart Leu-1\/T1 (CD5)?","volume":"84","author":"Huang","year":"1987","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023032406175973400_jlb10304-cit-0015","doi-asserted-by":"crossref","first-page":"4943","DOI":"10.4049\/jimmunol.160.10.4943","article-title":"The structure of avian CD5 implies a conserved function?","volume":"160","author":"Koskinen","year":"1998","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0016","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/1471-2199-2-5","article-title":"The regulation of CD5 expression in murine T cells?","volume":"2","author":"Tung","year":"2001","journal-title":"BMC Mol Biol"},{"key":"2023032406175973400_jlb10304-cit-0017","doi-asserted-by":"crossref","first-page":"3898","DOI":"10.1073\/pnas.0308764101","article-title":"The E47 transcription factor negatively regulates CD5 expression during thymocyte development?","volume":"101","author":"Yang","year":"2004","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023032406175973400_jlb10304-cit-0018","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1038\/icb.2009.109","article-title":"Increased antigen responsiveness of naive CD8 T cells exposed to IL-7 and IL-21 is associated with decreased CD5 expression?","volume":"88","author":"Gagnon","year":"2010","journal-title":"Immunol Cell Biol"},{"key":"2023032406175973400_jlb10304-cit-0019","doi-asserted-by":"crossref","first-page":"662","DOI":"10.1038\/351662a0","article-title":"The B-cell surface protein CD72\/Lyb-2 is the ligand for CD5?","volume":"351","author":"Van de Velde","year":"1991","journal-title":"Nature"},{"key":"2023032406175973400_jlb10304-cit-0020","doi-asserted-by":"crossref","first-page":"1279","DOI":"10.1084\/jem.184.4.1279","article-title":"CD5 is a potential selecting ligand for B cell surface immunoglobulin framework region sequences?","volume":"184","author":"Pospisil","year":"1996","journal-title":"J Exp Med"},{"key":"2023032406175973400_jlb10304-cit-0021","doi-asserted-by":"crossref","first-page":"6068","DOI":"10.4049\/jimmunol.0903823","article-title":"A ligand for CD5 is CD5?","volume":"185","author":"Brown","year":"2010","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0022","doi-asserted-by":"crossref","first-page":"1506","DOI":"10.1073\/pnas.0805846106","article-title":"The CD5 ectodomain interacts with conserved fungal cell wall components and protects from zymosan-induced septic shock-like syndrome?","volume":"106","author":"Vera","year":"2009","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2023032406175973400_jlb10304-cit-0023","doi-asserted-by":"crossref","first-page":"3723","DOI":"10.1128\/JVI.06956-11","article-title":"Hepatitis C virus infection of human T lymphocytes is mediated by CD5?","volume":"86","author":"Sarhan","year":"2012","journal-title":"J Virol"},{"key":"2023032406175973400_jlb10304-cit-0024","doi-asserted-by":"crossref","first-page":"6022","DOI":"10.4049\/jimmunol.161.11.6022","article-title":"Human CD5 signaling and constitutive phosphorylation of C-terminal serine residues by casein kinase II?","volume":"161","author":"Calvo","year":"1998","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0025","doi-asserted-by":"crossref","first-page":"19183","DOI":"10.1074\/jbc.273.30.19183","article-title":"Regulation of casein kinase 2 by direct interaction with cell surface receptor CD5?","volume":"273","author":"Raman","year":"1998","journal-title":"J Biol Chem"},{"key":"2023032406175973400_jlb10304-cit-0026","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1093\/intimm\/13.2.149","article-title":"Determination of the tyrosine phosphorylation sites in the T cell transmembrane glycoprotein CD5?","volume":"13","author":"Dennehy","year":"2001","journal-title":"Int 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Immunol"},{"key":"2023032406175973400_jlb10304-cit-0030","doi-asserted-by":"crossref","first-page":"1816","DOI":"10.4049\/jimmunol.137.6.1816","article-title":"Monoclonal antibodies to the CD5 antigen can provide the necessary second signal for activation of isolated resting T cells by solid-phase-bound OKT3?","volume":"137","author":"Ceuppens","year":"1986","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0031","doi-asserted-by":"crossref","first-page":"47","DOI":"10.4049\/jimmunol.146.1.47","article-title":"Induction of human T cell proliferation by a monoclonal antibody to CD5?","volume":"146","author":"Spertini","year":"1991","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0032","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.4049\/jimmunol.148.5.1287","article-title":"Intracellular events involved in CD5-induced human T cell activation and proliferation?","volume":"148","author":"Alberola-Ila","year":"1992","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0033","doi-asserted-by":"crossref","first-page":"4307","DOI":"10.4049\/jimmunol.159.9.4307","article-title":"The cytoplasmic domain of CD5 mediates both TCR\/CD3-dependent and -independent diacylglycerol production?","volume":"159","author":"Simarro","year":"1997","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0034","doi-asserted-by":"crossref","first-page":"5149","DOI":"10.4049\/jimmunol.162.9.5149","article-title":"Signaling through CD5 involves acidic sphingomyelinase, protein kinase C-zeta, mitogen-activated protein kinase kinase, and c-Jun NH2-terminal kinase?","volume":"162","author":"Simarro","year":"1999","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0035","doi-asserted-by":"crossref","first-page":"1678","DOI":"10.1002\/eji.1830240733","article-title":"Lymphocyte populations and immune responses in CD5-deficient 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Immunol"},{"key":"2023032406175973400_jlb10304-cit-0039","doi-asserted-by":"crossref","first-page":"2903","DOI":"10.1128\/MCB.19.4.2903","article-title":"CD5 negatively regulates the T-cell antigen receptor signal transduction pathway: involvement of SH2-containing phosphotyrosine phosphatase SHP-1?","volume":"19","author":"Perez-Villar","year":"1999","journal-title":"Mol Cell Biol"},{"key":"2023032406175973400_jlb10304-cit-0040","doi-asserted-by":"crossref","first-page":"45","DOI":"10.3892\/ijmm.2016.2592","article-title":"CD5-mediated inhibition of TCR signaling proceeds normally in the absence of SHP-1?","volume":"38","author":"Dong","year":"2016","journal-title":"Int J Mol Med"},{"key":"2023032406175973400_jlb10304-cit-0041","doi-asserted-by":"crossref","first-page":"679","DOI":"10.1002\/eji.1830270316","article-title":"Thymocyte activation induces the association of phosphatidylinositol 3-kinase and pp120 with CD5?","volume":"27","author":"Dennehy","year":"1997","journal-title":"Eur J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0042","doi-asserted-by":"crossref","first-page":"1617","DOI":"10.1002\/(SICI)1521-4141(199805)28:05<1617::AID-IMMU1617>3.0.CO;2-7","article-title":"Thymocyte activation induces the association of the proto-oncoprotein c-cbl and ras GTPase-activating protein with CD5?","volume":"28","author":"Dennehy","year":"1998","journal-title":"Eur J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0043","doi-asserted-by":"crossref","first-page":"876","DOI":"10.15252\/msb.20166837","article-title":"Co-recruitment analysis of the CBL and CBLB signalosomes in primary T cells identifies CD5 as a key regulator of TCR-induced ubiquitylation?","volume":"12","author":"Voisinne","year":"2016","journal-title":"Mol Syst Biol"},{"key":"2023032406175973400_jlb10304-cit-0044","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.bbrc.2013.01.086","article-title":"The carboxy-terminal region of CD5 is required for c-CBL mediated TCR signaling downmodulation in thymocytes?","volume":"432","author":"Roa","year":"2013","journal-title":"Biochem Biophys Res Commun"},{"key":"2023032406175973400_jlb10304-cit-0045","doi-asserted-by":"crossref","first-page":"5817","DOI":"10.4049\/jimmunol.161.11.5817","article-title":"Differential CD5-dependent regulation of CD5-associated CK2 activity in mature and immature T cells: implication on TCR\/CD3-mediated activation?","volume":"161","author":"Raman","year":"1998","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0046","doi-asserted-by":"crossref","first-page":"8542","DOI":"10.4049\/jimmunol.177.12.8542","article-title":"CD5-CK2 binding\/activation-deficient mice are resistant to experimental autoimmune encephalomyelitis: protection is associated with diminished populations of IL-17-expressing T cells in the central nervous system?","volume":"177","author":"Axtell","year":"2006","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0047","doi-asserted-by":"crossref","first-page":"7537","DOI":"10.4049\/jimmunol.1002173","article-title":"A conserved enhancer element differentially regulates developmental expression of CD5 in B and T cells?","volume":"185","author":"Berland","year":"2010","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0048","doi-asserted-by":"crossref","first-page":"2781","DOI":"10.1182\/blood-2005-02-0597","article-title":"An alternative exon 1 of the CD5 gene regulates CD5 expression in human B lymphocytes?","volume":"106","author":"Renaudineau","year":"2005","journal-title":"Blood"},{"key":"2023032406175973400_jlb10304-cit-0049","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1002\/eji.1830240406","article-title":"CD5 is associated with the human B cell antigen receptor complex?","volume":"24","author":"Lankester","year":"1994","journal-title":"Eur J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0050","doi-asserted-by":"crossref","first-page":"1906","DOI":"10.1126\/science.274.5294.1906","article-title":"CD5-mediated negative regulation of antigen receptor-induced growth signals in B-1 B cells?","volume":"274","author":"Bikah","year":"1996","journal-title":"Science"},{"key":"2023032406175973400_jlb10304-cit-0051","doi-asserted-by":"crossref","first-page":"548","DOI":"10.1074\/jbc.275.1.548","article-title":"The pseudo-immunoreceptor tyrosine-based activation motif of CD5 mediates its inhibitory action on B-cell receptor signaling?","volume":"275","author":"Gary-Gouy","year":"2000","journal-title":"J Biol Chem"},{"key":"2023032406175973400_jlb10304-cit-0052","doi-asserted-by":"crossref","first-page":"232","DOI":"10.4049\/jimmunol.168.1.232","article-title":"CD5-negative regulation of B cell receptor signaling pathways originates from tyrosine residue Y429 outside an immunoreceptor tyrosine-based inhibitory motif?","volume":"168","author":"Gary-Gouy","year":"2002","journal-title":"J Immunol"},{"key":"2023032406175973400_jlb10304-cit-0053","doi-asserted-by":"crossref","first-page":"1417","DOI":"10.1093\/intimm\/12.10.1417","article-title":"Negative regulation of B cell receptor-mediated signaling in B-1 cells through CD5 and Ly49 co-receptors via Lyn kinase activity?","volume":"12","author":"Ochi","year":"2000","journal-title":"Int Immunol"},{"key":"2023032406175973400_jlb10304-cit-0054","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1016\/j.immuni.2016.04.003","article-title":"CD5 binds to Interleukin-6 and induces a feed-forward loop with the transcription factor STAT3 in B cells to promote cancer?","volume":"44","author":"Zhang","year":"2016","journal-title":"Immunity"},{"key":"2023032406175973400_jlb10304-cit-0055","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1016\/j.bbrc.2017.01.174","article-title":"Trans-signaling of interleukin-6 (IL-6) is mediated by the soluble IL-6 receptor, but not by soluble CD5?","volume":"484","author":"Aparicio-Siegmund","year":"2017","journal-title":"Biochem Biophys Res Commun"},{"key":"2023032406175973400_jlb10304-cit-0056","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1182\/blood-2007-08-110130","article-title":"Scan of 977 nonsynonymous SNPs in CLL4 trial patients for the identification of genetic variants influencing prognosis?","volume":"111","author":"Sellick","year":"2008","journal-title":"Blood"},{"key":"2023032406175973400_jlb10304-cit-0057","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1111\/bjh.14037","article-title":"Impact of the functional CD5 polymorphism A471V on the response of chronic lymphocytic leukaemia to conventional chemotherapy regimens?","volume":"177","author":"Delgado","year":"2017","journal-title":"Br J Haematol"},{"key":"2023032406175973400_jlb10304-cit-0058","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.1002\/ijc.30184","article-title":"Inherited functional variants of the lymphocyte receptor CD5 influence melanoma survival?","volume":"139","author":"Potrony","year":"2016","journal-title":"Int J Cancer"},{"key":"2023032406175973400_jlb10304-cit-0059","doi-asserted-by":"crossref","first-page":"e113090","DOI":"10.1371\/journal.pone.0113090","article-title":"Analysis of ancestral and functionally relevant CD5 variants in systemic lupus erythematosus patients?","volume":"9","author":"Cenit","year":"2014","journal-title":"PLoS One"},{"key":"2023032406175973400_jlb10304-cit-0060","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1038\/sj.icb.7100141","article-title":"What do we know about the mechanisms of elimination of autoreactive T and 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