{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T03:49:43Z","timestamp":1790653783900,"version":"4.1.0"},"reference-count":167,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2024,7,4]],"date-time":"2024-07-04T00:00:00Z","timestamp":1720051200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Mark Terasaki, the first son of the late Paul Ichiro Terasaki of the Terasaki Foundation Laboratory (TFL)"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["CIMB"],"abstract":"<jats:p>Human leukocyte antigens (HLAs) are polymorphic glycoproteins expressed on the cell surface of nucleated cells and consist of two classes, HLA class I and HLA class II. In contrast, in mice, these molecules, known as H-2, are expressed on both nucleated cells and erythrocytes. HLA-I molecules (Face-1) are heterodimers consisting of a polypeptide heavy chain (HC) and a light chain, B2-microglobulin (B2m). The heterodimers bind to antigenic peptides and present them to the T-cell receptors of CD8+ cytotoxic T lymphocytes. The HCs can also independently emerge on the cell surface as B2m-free HC monomers without peptides (Face-2). Early investigators suggested that the occurrence of B2m-free HCs on the cell surface resulted from the dissociation of B2m from Face-1. However, others documented the independent emergence of B2m-free HCs (Face-2) from the endoplasmic reticulum (ER) to the cell surface. The clustering of such HC molecules on either the cell surface or on exosomes resulted in the dimerization of B2m-free HCs to form homodimers (if the same allele, designated as Face-3) or heterodimers (if different alleles, designated as Face-4). Face-2 occurs at low levels on the cell surface of several normal cells but is upregulated on immune cells upon activation by proinflammatory cytokines and other agents such as anti-CD3 antibodies, phytohemagglutinin, and phorbol myristate acetate. Their density on the cell surface remains high as long as the cells remain activated. After activation-induced upregulation, Face-2 molecules undergo homo- and heterodimerization (Face-3 and Face-4). Observations made on the structural patterns of HCs and their dimerization in sharks, fishes, and tetrapod species suggest that the formation of B2m-free HC monomers and dimers is a recapitalization of a phylogenetically conserved event, befitting the term Proto-HLA for the B2m-free HCs. Spontaneous arthritis occurs in HLA-B27+ mice lacking B2m (HLA-B27+ B2m\u2212\/\u2212) but not in HLA-B27+ B2m+\/+ mice. Anti-HC-specific monoclonal antibodies (mAbs) delay disease development. Some HLA-I polyreactive mAbs (MEM series) used for immunostaining confirm the existence of B2m-free variants in several cancer cells. The conformational alterations that occur in the B2m-free HCs enable them to interact with several inhibitory and activating receptors of cellular components of the innate (natural killer (NK) cells) and adaptive (T and B cells) immune systems. The NK cells express killer immunoglobulin-like receptors (KIRs), whereas leukocytes (T and B lymphocytes, monocytes\/macrophages, and dendritic cells) express leukocyte immunoglobulin-like receptors (LILRs). The KIRs and LILRs include activating and inhibitory members within their respective groups. This review focuses on the interaction of KIRs and LILRs with B2m-free HC monomers and dimers in patients with spondylarthritis. Several investigations reveal that the conformational alterations occurring in the alpha-1 and alpha-2 domains of B2m-free HCs may facilitate immunomodulation by their interaction with KIR and LILR receptors. This opens new avenues to immunotherapy of autoimmune diseases and even human cancers that express B2m-free HCs.<\/jats:p>","DOI":"10.3390\/cimb46070416","type":"journal-article","created":{"date-parts":[[2024,7,4]],"date-time":"2024-07-04T09:50:32Z","timestamp":1720086632000},"page":"6961-6985","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Conformational Alterations of the Cell Surface of Monomeric and Dimeric \u03b22m-Free HLA-I (Proto-HLA) May Enable Novel Immune Functions in Health and Disease"],"prefix":"10.3390","volume":"46","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5731-4129","authenticated-orcid":false,"given":"Mepur H.","family":"Ravindranath","sequence":"first","affiliation":[{"name":"Department of Hematology and Oncology, Children\u2019s Hospital, Los Angeles, CA 90027, USA"},{"name":"Terasaki Foundation Laboratory, Santa Monica, CA 90064, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Narendranath M.","family":"Ravindranath","sequence":"additional","affiliation":[{"name":"Norris Dental Science Center, Herman Ostrow School of Dentistry, University of Southern California, Los Angeles, CA 90089, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carly J.","family":"Amato-Menker","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology and Cell Biology, School of Medicine, West Virginia University, Morgantown, WV 26506, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fatiha El","family":"Hilali","sequence":"additional","affiliation":[{"name":"Medico-Surgical, Biomedicine and Infectiology Research Laboratory, The Faculty of Medicine and Pharmacy of Laayoune & Agadir, Ibnou Zohr University, Agadir 80000, Morocco"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3161-5820","authenticated-orcid":false,"given":"Edward J.","family":"Filippone","sequence":"additional","affiliation":[{"name":"Division of Nephrology, Department of Medicine, Sidney Kimmel Medical College, Thomas Jefferson University, Philadelphia, PA 19145, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2024,7,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2123","DOI":"10.1073\/pnas.71.5.2123","article-title":"Immunological identity of the small subunit of HL-A antigens and beta2-microglobulin and its turnover on the cell membrane","volume":"71","author":"Cresswell","year":"1974","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"2481","DOI":"10.1073\/pnas.73.7.2481","article-title":"Detergent-soluble HLA antigens contain a hydrophilic region at the COOH-terminus and a penultimate hydrophobic region","volume":"73","author":"Springer","year":"1976","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"7555","DOI":"10.1016\/S0021-9258(17)41003-9","article-title":"Carbohydrate moiety of HLA antigens. Antigenic properties and amino acid sequences around the site of glycosylation","volume":"252","author":"Parham","year":"1977","journal-title":"J. Biol. Chem."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Ravindranath, M.H., Ravindranath, N.M., Selvan, S.R., Hilali, F.E., Amato-Menker, C.J., and Filippone, E.J. (2023). Cell Surface B2m-Free Human Leukocyte Antigen (HLA) Monomers and Dimers: Are They Neo-HLA Class and Proto-HLA?. Biomolecules, 13.","DOI":"10.3390\/biom13081178"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1038\/254267a0","article-title":"The B2-microglobulin gene is on chromosome 15 and not in the HLA region","volume":"254","author":"Goodfellow","year":"1975","journal-title":"Nature"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1111\/j.1399-0039.1978.tb01233.x","article-title":"The genetic control of HLA-A and B antigens in somatic cell hybrids: Requirement for beta2 microglobulin","volume":"11","author":"Jones","year":"1978","journal-title":"Tissue Antigens"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"7447","DOI":"10.1073\/pnas.83.19.7447","article-title":"Beta 2-microglobulin is not required for cell surface expression of the murine class I histocompatibility antigen H-2Db or of a truncated H-2Db","volume":"83","author":"Allen","year":"1986","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1016\/0092-8674(82)90182-9","article-title":"Structure of wild-type and mutant mouse beta 2-microglobulin genes","volume":"29","author":"Parnes","year":"1982","journal-title":"Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1073\/pnas.76.5.2273","article-title":"Cell-free translation of the mRNAs for the heavy and light chains of HLA-A and HLA-B antigens","volume":"76","author":"Ploegh","year":"1979","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1084\/jem.160.1.317","article-title":"Expression of H-2Db on the cell surface in the absence of detectable beta 2 microglobulin","volume":"160","author":"Potter","year":"1984","journal-title":"J. Exp. Med."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"2950","DOI":"10.1073\/pnas.82.9.2950","article-title":"Molecular analysis of an EL4 cell line that expresses H-2Db but not H-2Kb or beta 2-microglobulin","volume":"82","author":"Potter","year":"1985","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1002\/(SICI)1521-4141(199801)28:01<370::AID-IMMU370>3.0.CO;2-W","article-title":"Beta2-microglobulin-deficient NK cells show increased sensitivity to MHC class I-mediated inhibition, but self tolerance does not depend upon target cell expression of H-2Kb and Db heavy chains","volume":"28","author":"Glas","year":"1998","journal-title":"Eur. J. Immunol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3522","DOI":"10.4049\/jimmunol.140.10.3522","article-title":"Studies of two antigenic forms of Ld with disparate beta 2-microglobulin (beta 2m) associations suggest that beta 2m facilitate the folding of the alpha 1 and alpha 2 domains during de novo synthesis","volume":"140","author":"Hansen","year":"1988","journal-title":"J. Immunol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1016\/0092-8674(91)90093-E","article-title":"Dissociation of beta 2-microglobulin leads to the accumulation of a substantial pool of inactive class I MHC heavy chains on the cell surface","volume":"65","author":"Rock","year":"1991","journal-title":"Cell"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2751","DOI":"10.4049\/jimmunol.142.8.2751","article-title":"The conformation of Ld induced by beta 2-microglobulin is fixed during de novo synthesis and irreversible by exchange or dissociation","volume":"142","author":"Myer","year":"1989","journal-title":"J. Immunol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2796","DOI":"10.4049\/jimmunol.142.8.2796","article-title":"Role of beta 2-microglobulin in the intracellular transport and surface expression of murine class I histocompatibility molecules","volume":"142","author":"Williams","year":"1989","journal-title":"J. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1016\/0092-8674(79)90210-1","article-title":"Assembly and maturation of HLA-A and HLA-B antigens in vivo","volume":"18","author":"Krangel","year":"1979","journal-title":"Cell"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/0092-8674(78)90296-9","article-title":"Production of monoclonal antibodies to group A erythrocytes, HLA and other human cell surface antigens-new tools for genetic analysis","volume":"14","author":"Barnstable","year":"1978","journal-title":"Cell"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"342","DOI":"10.4049\/jimmunol.123.1.342","article-title":"Use of a monoclonal antibody (W6\/32) in structural studies of HLA-A,B,C, antigens","volume":"123","author":"Parham","year":"1979","journal-title":"J. Immunol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1073\/pnas.83.2.432","article-title":"Unusual association of beta 2-microglobulin with certain class I heavy chains of the murine major histocompatibility complex","volume":"83","author":"Bushkin","year":"1986","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1084\/jem.164.2.458","article-title":"A new HLA-linked T cell membrane molecule, related to the beta chain of the clonotypic receptor, is associated with T3","volume":"164","author":"Bushkin","year":"1986","journal-title":"J. Exp. Med."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3985","DOI":"10.1073\/pnas.85.11.3985","article-title":"Physical association between the CD8 and HLA class I molecules on the surface of activated human T lymphocytes","volume":"85","author":"Bushkin","year":"1988","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1431","DOI":"10.1084\/jem.171.5.1431","article-title":"Activated human T lymphocytes express MHC class I heavy chains not associated with beta 2-microglobulin","volume":"171","author":"Schnabl","year":"1990","journal-title":"J. Exp. Med."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/0008-8749(92)90272-Q","article-title":"The origin and fate of beta 2m-free MHC class I molecules induced on activated T cells","volume":"142","author":"Demaria","year":"1992","journal-title":"Cell Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1007\/BF02592285","article-title":"CD8 and beta 2-microglobulin-free MHC class I molecules in T cell immunoregulation","volume":"23","author":"Demaria","year":"1993","journal-title":"Int. J. Clin. Lab. Res."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1007\/s002510100353","article-title":"The epitope recognized by pan-HLA class I-reactive monoclonal antibody W6\/32 and its relationship to unusual stability of the HLA-B27\/beta2-microglobulin complex","volume":"53","author":"Tran","year":"2001","journal-title":"Immunogenetics"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1080\/21548331.1979.11707628","article-title":"Seronegative spondylarthropathies","volume":"14","author":"Bluestone","year":"1979","journal-title":"Hosp. Pract."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Bluestone, R. (1979). Diagnosis of rheumatic disease. 2. Laboratory tests. Postgrad. Med., 65.","DOI":"10.1080\/00325481.1979.11715082"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1084\/jem.182.4.1153","article-title":"Spontaneous inflammatory arthritis in HLA-B27 transgenic mice lacking beta 2-microglobulin: A model of human spondyloarthropathies","volume":"182","author":"Khare","year":"1995","journal-title":"J. Exp. Med."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2746","DOI":"10.1172\/JCI119100","article-title":"HLA-B27 heavy chains contribute to spontaneous inflammatory disease in B27\/human beta2-microglobulin (beta2m) double transgenic mice with disrupted mouse beta2m","volume":"98","author":"Khare","year":"1996","journal-title":"J. Clin. Investig."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2299","DOI":"10.4049\/jimmunol.137.7.2299","article-title":"Monoclonal antibodies raised against denatured HLA-B locus heavy chains permit biochemical characterization of certain HLA-C locus products","volume":"137","author":"Stam","year":"1986","journal-title":"J. Immunol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1918","DOI":"10.4049\/jimmunol.171.4.1918","article-title":"Beta 2-microglobulin-free HLA class I heavy chain epitope mimicry by monoclonal antibody HC-10-specific peptide","volume":"171","author":"Perosa","year":"2003","journal-title":"J. Immunol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"3139","DOI":"10.1002\/art.34538","article-title":"Inhibiting HLA-B27 homodimer-driven immune cell inflammation in spondylarthritis","volume":"64","author":"Payeli","year":"2012","journal-title":"Arthritis Rheum."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"763","DOI":"10.1136\/annrheumdis-2012-203080","article-title":"Expression of aberrant HLA-B27 molecules is dependent on B27 dosage and peptide supply","volume":"73","author":"McHugh","year":"2014","journal-title":"Ann Rheum Dis."},{"key":"ref_35","first-page":"966","article-title":"Free HLA class I heavy chain-carrying monocytes\u2014A potential role in the pathogenesis of spondyloarthropathies","volume":"29","author":"Tsai","year":"2002","journal-title":"J. Rheumatol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1111\/j.1365-2133.2004.05890.x","article-title":"Psoriatic patients with arthropathy show significant expression of free HLA class I heavy chains on circulating monocytes: A potential role in the pathogenesis of psoriatic arthropathy","volume":"151","author":"Lan","year":"2004","journal-title":"Br. J. Dermatol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1338","DOI":"10.1093\/rheumatology\/kel305","article-title":"Consistent patterns of expression of HLA class I free heavy chains in healthy individuals and raised expression in spondyloarthropathy patients point to physiological and pathological roles","volume":"45","author":"Raine","year":"2006","journal-title":"Rheumatology"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"R100","DOI":"10.1186\/ar2492","article-title":"Induction of HLA-B27 heavy chain homodimer formation after activation in dendritic cells","volume":"10","author":"Santos","year":"2008","journal-title":"Arthritis Res. Ther."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"16180","DOI":"10.1073\/pnas.212643199","article-title":"Disulfide bond-mediated dimerization of HLA-G on the cell surface","volume":"99","author":"Boyson","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1016\/S0161-5890(98)00077-7","article-title":"The monoclonal antibody HCA2 recognises a broadly shared epitope on selected classical as well as several non-classical HLA class I molecules","volume":"35","author":"Seitz","year":"1998","journal-title":"Mol. Immunol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1935","DOI":"10.4049\/jimmunol.1000424","article-title":"Antibodies to HLA-E in non-alloimmunized Males: Pattern of HLA-Ia-reactivity of anti\u2013HLA-E\u2013positive sera","volume":"185","author":"Ravindranath","year":"2010","journal-title":"J. Immunol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"6199","DOI":"10.4049\/jimmunol.1000078","article-title":"HLA-F complex without peptide binds to MHC class I protein in the open conformer form","volume":"184","author":"Goodridge","year":"2010","journal-title":"J. Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"jcs259489","DOI":"10.1242\/jcs.259498","article-title":"Dissociation of \u03b22m from MHC class I triggers formation of noncovalent transient heavy chain dimers","volume":"135","author":"Dirscherl","year":"2022","journal-title":"J. Cell Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1002\/eji.1830150915","article-title":"A third polypeptide associated with heavy and light chain subunits of class I HLA antigens in immune interferon-treated human melanoma cells","volume":"15","author":"Giacomini","year":"1985","journal-title":"Eur. J. Immunol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1111\/j.1399-0039.1997.tb02913.x","article-title":"HLA-C heavy chains free of beta2-microglobulin: Distribution in normal tissues and neoplastic lesions of non-lymphoid origin and interferon-y responsiveness","volume":"50","author":"Giacomini","year":"1997","journal-title":"Tissue Antigens"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1111\/j.1365-3083.1993.tb01680.x","article-title":"Expression of beta 2m-free HLA class I heavy chains in neuroblastoma cell lines","volume":"37","author":"Marozzi","year":"1993","journal-title":"Scand. J. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/S0198-8859(96)00256-X","article-title":"Conformation and surface expression of free HLA-CW1 heavy chains in the absence of beta 2-microglobulin","volume":"53","author":"Martayan","year":"1997","journal-title":"Hum. Immunol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1016\/j.molimm.2009.10.024","article-title":"HLA-E monoclonal antibodies recognize shared peptide sequences on classical HLA class Ia: Relevance to human natural HLA antibodies","volume":"47","author":"Ravindranath","year":"2010","journal-title":"Mol. Immunol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/j.molimm.2010.09.011","article-title":"Anti-HLAE mAb 3D12 mimics MEM-E\/02 in binding to HLA-B and HLA-C alleles: Web-tools validate the immunogenic epitopes of HLA-E recognized by the antibodies","volume":"48","author":"Ravindranath","year":"2011","journal-title":"Mol. Immunol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/S0198-8859(02)00821-2","article-title":"Characterization of monoclonal antibodies recognizing HLA-G or HLA-E: New tools to analyze the expression of nonclassical HLA class I molecules","volume":"64","author":"Menier","year":"2003","journal-title":"Hum. Immunol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"4951","DOI":"10.4049\/jimmunol.160.10.4951","article-title":"HLA-Esurface expression depends on binding of TAP-dependent peptides derived from certain HLAClass I signal sequences","volume":"160","author":"Lee","year":"1998","journal-title":"J. Immunol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1007\/s12026-018-9027-2","article-title":"Significance of the intraindividual variability of HLA IgG antibodies in renal disease patients observed with different beadsets monitored with two different secondary antibodies on a Luminex platform","volume":"66","author":"Ravindranath","year":"2018","journal-title":"Immunol. Res."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Ravindranath, M.H., Ravindranath, N.M., and Amato-Menker, C.J. (2021). Luminex Multiplex Bead Assay Monitoring HLA IgG Antibodies in Sensitized Pre- and Post-transplant Patients: Clonality of the Detection Antibody Impacts Specificity and Sensitivity. Appl. Sci., 11.","DOI":"10.3390\/app11146430"},{"key":"ref_54","first-page":"1","article-title":"HLA-E restricted monoclonal antibodies: Therapeutic potential as a double-edged sword against tumor progression","volume":"3","author":"Ravindranath","year":"2017","journal-title":"Internal Med. Rev."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1558","DOI":"10.1002\/ijc.28484","article-title":"Gastric cancer progression may involve a shift in HLA-E profile from an intact heterodimer to \u03b22-microglobulin-free monomer","volume":"134","author":"Sasaki","year":"2014","journal-title":"Int. J. Cancer"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"3100","DOI":"10.4049\/jimmunol.177.5.3100","article-title":"Expression and release of HLA-E by melanoma cells and melanocytes: Potential impact on the response of cytotoxic effector cells","volume":"177","author":"Corvaisier","year":"2006","journal-title":"J. Immunol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1016\/j.humimm.2015.12.001","article-title":"Relevance of HLA-G, HLA-E and IL-10 expression in lip carcinogenesis","volume":"77","author":"Oliveira","year":"2016","journal-title":"Hum. Immunol."},{"key":"ref_58","first-page":"1487","article-title":"Expression of the nonclassical HLA-G and HLA-E molecules in laryngeal lesions as biomarkers of tumor invasiveness","volume":"26","author":"Silva","year":"2011","journal-title":"Histol. Histopathol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1002\/ijc.28713","article-title":"Alterations in classical and nonclassical HLA expression in recurrent and progressive HPV-induced usual vulvar intraepithelial neoplasia and implications for immunotherapy","volume":"135","author":"Tummers","year":"2014","journal-title":"Int. J. Cancer."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1245","DOI":"10.1016\/j.juro.2014.11.057","article-title":"Classic and nonclassic HLA class I expression in penile cancer and relation to HPV status and clinical outcome","volume":"193","author":"Djajadiningrat","year":"2015","journal-title":"J. Urol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.jneuroim.2007.07.002","article-title":"Elevated HLA-Elevels in human glioblastomas but not in grade I to III astrocytomas correlate with infiltrating CD8+ cells","volume":"189","author":"Mittelbronn","year":"2007","journal-title":"J. Neuroimmunol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/j.jneuroim.2010.01.014","article-title":"Production of immune-modulatory nonclassical molecules HLA-G and HLA-E by tumor infiltrating ameboid microglia\/macrophages in glioblastomas: A role in innate immunity?","volume":"220","author":"Kren","year":"2010","journal-title":"J. Neuroimmunol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1111\/j.1440-1789.2010.01149.x","article-title":"Expression of immune-modulatory molecules HLA-G and HLA-E by tumor cells in glioblastomas: An unexpected prognostic significance?","volume":"31","author":"Kren","year":"2011","journal-title":"Neuropathology"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"e188","DOI":"10.1016\/j.oooo.2016.12.002","article-title":"Evaluation of HLA-G, HLA-E, and PD-L1 proteins in oral osteosarcomas","volume":"123","author":"Jham","year":"2017","journal-title":"Oral Surg. Oral Med. Oral Pathol. Oral Radiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.archoralbio.2017.07.004","article-title":"Immunohistochemical investigations on the expression of programmed cell death ligand 1, human leukocyte antigens G and E, and granzyme B in intraoral mucoepidermoid carcinoma","volume":"83","author":"Mosconi","year":"2017","journal-title":"Arch. Oral. Biol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Reimers, M.S., Engels, C.C., Putter, H., Morreau, H., Liefers, G.J., van de Velde, C.J., and Kuppen, P.J. (2014). Prognostic value of HLA class I, HLA-E, HLA-G and Tregs in rectal cancer: A retrospective cohort study. BMC Cancer, 14.","DOI":"10.1186\/1471-2407-14-486"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1186\/1479-5876-9-184","article-title":"High expression of HLA-E in colorectal carcinoma is associated with a favorable prognosis","volume":"9","author":"Benevolo","year":"2011","journal-title":"J. Transl. Med."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1038\/bjc.2013.696","article-title":"Combined analysis of HLA class I, HLA-E and HLA-G predicts prognosis in colon cancer patients","volume":"110","author":"Zeestraten","year":"2014","journal-title":"Br. J. Cancer."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.cellimm.2014.10.003","article-title":"Predictive value of HLA-G and HLA-E in the prognosis of colorectal cancer patients","volume":"293","author":"Guo","year":"2015","journal-title":"Cell. Immunol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"3379","DOI":"10.3892\/ol.2017.5891","article-title":"Loss of Fas expression and high expression of HLA-E promoting the immune escape of early colorectal cancer cells","volume":"13","author":"Huang","year":"2017","journal-title":"Oncol. Lett."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"371","DOI":"10.4149\/neo_2011_05_371","article-title":"Expression of the nonclassical HLA class I and MICA\/B molecules in human hepatocellular carcinoma","volume":"58","author":"Chen","year":"2011","journal-title":"Neoplasma"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"3477","DOI":"10.18632\/oncotarget.6506","article-title":"The positive prognostic effect of stromal CD8+ tumor-infiltrating T cells is restrained by the expression of HLA-E in non-small cell lung carcinoma","volume":"7","author":"Fiocco","year":"2016","journal-title":"Oncotarget"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"7452","DOI":"10.4049\/jimmunol.1002629","article-title":"HLA-E and HLA-G expression in classical HLA class I-negative tumors is of prognostic value for clinical outcome of early breast cancer patients","volume":"185","author":"Sajet","year":"2010","journal-title":"J. Immunol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"250435","DOI":"10.1155\/2013\/250435","article-title":"Expression of the Classical and Nonclassical HLA Molecules in Breast Cancer","volume":"2013","author":"Silva","year":"2013","journal-title":"Int. J. Breast Cancer"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"10656","DOI":"10.1073\/pnas.1100354108","article-title":"HLA-E expression by gynecological cancers restrains tumor-infiltrating CD8+ T lymphocytes","volume":"108","author":"Gooden","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.ejogrb.2008.06.010","article-title":"Classical and non-classical HLA molecules and p16 (INK4a) expression in precursors lesions and invasive cervical cancer","volume":"141","author":"Rabreau","year":"2008","journal-title":"Eur. J. Obstet. Gynecol. Reprod. Biol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1186\/1479-5876-10-184","article-title":"HLA-E expression in cervical adenocarcinomas: Association with improved long-term survival","volume":"10","author":"Spaans","year":"2012","journal-title":"J. Transl. Med."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1186\/s40425-016-0184-3","article-title":"Classical and nonclassical HLA class I aberrations in primary cervical squamous- and adenocarcinomas and paired lymph node metastases","volume":"4","author":"Ferns","year":"2016","journal-title":"J. Immunother. Cancer"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"e1052213","DOI":"10.1080\/2162402X.2015.1052213","article-title":"Non-classical HLA-class I expression in serous ovarian carcinoma: Correlation with the HLA-genotype, tumor infiltrating immune cells and prognosis","volume":"5","author":"Andersson","year":"2015","journal-title":"Oncoimmunology"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"522","DOI":"10.1111\/cas.12641","article-title":"Human leukocyte antigen-E alleles and expression in patients with serous ovarian cancer","volume":"106","author":"Zheng","year":"2015","journal-title":"Cancer Sci."},{"key":"ref_81","first-page":"CR638","article-title":"Expression pattern of HLA class I antigens in renal cell carcinoma and primary cell line cultures: Methodological implications for immunotherapy","volume":"15","author":"Hanak","year":"2009","journal-title":"Med. Sci. Monit."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1186\/1746-1596-7-58","article-title":"HLA-G and HLA-E specific mRNAs connote opposite prognostic significance in renal cell carcinoma","volume":"7","author":"Kren","year":"2012","journal-title":"Diagn. Pathol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1177\/039463201302600407","article-title":"Differential expression of immunemodulatory molecule HLA-E in nonneoplastic and neoplastic lesions of the thyroid","volume":"26","author":"Zanetti","year":"2013","journal-title":"Int. J. Immunopathol. Pharmacol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.prp.2011.11.004","article-title":"Multifunctional immune-modulatory protein HLA-E identified in classical Hodgkin lymphoma: Possible implications","volume":"208","author":"Kren","year":"2012","journal-title":"Pathol. Res. Pract."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1007\/s00251-002-0526-9","article-title":"Analysis of HLA-E expression in human tumors","volume":"54","author":"Pedrinaci","year":"2003","journal-title":"Immunogenetics"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.jneuroim.2012.05.010","article-title":"HLA-E contributes to an immune inhibitory phenotype of glioblastoma stem-like cells","volume":"250","author":"Wolpert","year":"2012","journal-title":"J. Neuroimmunol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1093\/jnen\/64.6.523","article-title":"HLA-E protects glioma cells from NKG2D mediated immune responses in vitro: Implications for immune escape in vivo","volume":"64","author":"Wischhusen","year":"2005","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"7465741","DOI":"10.1155\/2016\/7465741","article-title":"Soluble HLA-G and HLA-E Levels in Bone Marrow Plasma Samples Are Related to Disease Stage in Neuroblastoma Patients","volume":"2016","author":"Morandi","year":"2016","journal-title":"J. Immunol. Res."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"e1226720","DOI":"10.1080\/2162402X.2016.1226720","article-title":"Therapeutic CD94\/NKG2A blockade improves natural killer cell dysfunction in chronic lymphocytic leukemia","volume":"5","author":"McWilliams","year":"2016","journal-title":"Oncoimmunology"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1002\/cncr.30427","article-title":"HLA-E allelic genotype correlates with HLA-E plasma levels and predicts early progression in chronic lymphocytic leukemia","volume":"123","author":"Wagner","year":"2017","journal-title":"Cancer"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1093\/intimm\/dxn141","article-title":"Peptides with dual binding specificity for HLA-A2 and HLA-E are encoded by alternatively spliced isoforms of the antioxidant enzyme peroxiredoxin 5","volume":"21","author":"Sensi","year":"2009","journal-title":"Int. Immunol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"159","DOI":"10.4049\/jimmunol.151.1.159","article-title":"In vivo dimeric association of class I MHC heavy chains. Possible relationship to class I MHC heavy chain-beta 2-microglobulin dissociation","volume":"151","author":"Capps","year":"1993","journal-title":"J. Immunol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"21263","DOI":"10.1016\/S0021-9258(19)36919-4","article-title":"The cytoplasmic domain of the H-2Ld class I major histocompatibility complex molecule is differentially accessible to immunological and biochemical probes during transport to the cell surface","volume":"268","author":"Capps","year":"1993","journal-title":"J. Biol. Chem."},{"key":"ref_94","first-page":"2826","article-title":"Establishment and characterization of a mouse tumor cell line with irreversible downregulation of MHC class I molecules","volume":"42","author":"Lhotakova","year":"2019","journal-title":"Oncol. Rep."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"7182","DOI":"10.1021\/bi00146a022","article-title":"Self-association of class I major histocompatibility complex molecules in liposome and cell surface membranes","volume":"31","author":"Chakrabarti","year":"1992","journal-title":"Biochemistry"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"3353","DOI":"10.4049\/jimmunol.152.7.3353","article-title":"Clustering of class I HLA molecules on the surfaces of activated and transformed human cells","volume":"152","author":"Matko","year":"1994","journal-title":"J. Immunol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1007\/s002510050240","article-title":"Probing the stability of class I major histocompatibility complex (MHC) molecules on the surface of human cells","volume":"46","author":"Edidin","year":"1997","journal-title":"Immunogenetics"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1007\/s002510050596","article-title":"The role of HLA-B27 in spondyloarthritis","volume":"50","author":"Allen","year":"1999","journal-title":"Immunogenetics"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"1343","DOI":"10.4049\/jimmunol.171.3.1343","article-title":"Complexes of HLA-G protein on the cell surface are important for leukocyte Ig-like receptor-1 function","volume":"171","author":"Achdout","year":"2003","journal-title":"J. Immunol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"4866","DOI":"10.4049\/jimmunol.175.8.4866","article-title":"The CD85J\/leukocyte inhibitory receptor-1 distinguishes between conformed and beta 2-microglobulin-free HLA-G molecules","volume":"175","author":"Achdout","year":"2005","journal-title":"J. Immunol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"1122","DOI":"10.1073\/pnas.92.4.1122","article-title":"Structural hierarchy in the clustering of HLA class I molecules in the plasma membrane of human lymphoblastoid cells","volume":"92","author":"Dmjanovich","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1093\/intimm\/dxg042","article-title":"Class I HLA oligomerization at the surface of B cells is controlled by exogenous beta(2)-microglobulin: Implications in activation of cytotoxic T lymphocytes","volume":"15","author":"Jenei","year":"2003","journal-title":"Int. Immunol."},{"key":"ref_103","doi-asserted-by":"crossref","unstructured":"Kennedy, P.R., Barthen, C., Williamson, D.J., and Davies, D.M. (2019). HLA-B and HLA-C differ in their nanoscale organization at cell surfaces. Front. Immunol., 10.","DOI":"10.3389\/fimmu.2019.00061"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"6665","DOI":"10.4049\/jimmunol.163.12.6665","article-title":"Misfolding of HLA-B27 as a result of its B pocket suggests a novel mechanism for its role in susceptibility to spondyloarthropathies","volume":"163","author":"Mear","year":"1999","journal-title":"J. Immunol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1002\/eji.200323678","article-title":"Lymphoblastoid cells express HLA-B27 homodimers both intracellularly and at the cell surface following endosomal recycling","volume":"33","author":"Bird","year":"2003","journal-title":"Eur. J. Immunol."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"5110","DOI":"10.4049\/jimmunol.172.8.5110","article-title":"HLA-B27in transgenic rats forms disulfide-linked heavy chain oligomers and multimers that bind to the chaperone BiP","volume":"172","author":"Tran","year":"2004","journal-title":"J. Immunol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1053\/gast.2001.26263","article-title":"Intestinal epithelial cells secrete exosome-like vesicles","volume":"121","author":"Raposo","year":"2001","journal-title":"Gastroenterology"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"9412","DOI":"10.1016\/S0021-9258(18)48095-7","article-title":"Vesicle formation during reticulocyte maturation. Association of plasma membrane activities with released vesicles (exosomes)","volume":"262","author":"Johnstone","year":"1987","journal-title":"J. Biol. Chem."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"642","DOI":"10.1016\/j.mcn.2005.12.003","article-title":"Sadoul Exosomes are released by cultured cortical neurons","volume":"31","author":"Lachenal","year":"2006","journal-title":"Mol. Cell. Neurosci."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"2631","DOI":"10.1091\/mbc.8.12.2631","article-title":"Accumulation of major histocompatibility complex class II molecules in mast cell secretory granules and their release upon degranulation","volume":"8","author":"Raposo","year":"1997","journal-title":"Mol. Biol. Cell"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.1084\/jem.183.3.1161","article-title":"B lymphocytes secrete antigen-presenting vesicles","volume":"183","author":"Raposo","year":"1996","journal-title":"J. Exp. Med."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"5242","DOI":"10.4049\/jimmunol.181.8.5242","article-title":"Thymus exosomes-like particles induce regulatory T cells","volume":"181","author":"Wang","year":"2008","journal-title":"J. Immunol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1615\/CritRevImmunol.v25.i3.30","article-title":"Dendritic cell-derived exosomes as cell-free peptide-based vaccines","volume":"25","author":"Chaput","year":"2005","journal-title":"Crit. Rev. Immunol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"6440","DOI":"10.4049\/jimmunol.174.10.6440","article-title":"Exosomes derived from IL-10-treated dendritic cells can suppress inflammation and collagen-induced arthritis","volume":"174","author":"Kim","year":"2005","journal-title":"J. Immunol."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/1479-5876-3-10","article-title":"Zitvogel Vaccination of metastatic melanoma patients with autologous dendritic cell (DC) derived-exosomes: Results of thefirst phase I clinical trial","volume":"3","author":"Escudier","year":"2005","journal-title":"J. Transl. Med."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"376","DOI":"10.1111\/j.1582-4934.2006.tb00406.x","article-title":"Prospects for exosomes in immunotherapy of cancer","volume":"10","author":"Mignot","year":"2006","journal-title":"J. Cell Mol. Med."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"7864","DOI":"10.1158\/0008-5472.CAN-07-6538","article-title":"Senescence-associated exosome release from human prostate cancer cells","volume":"68","author":"Lehmann","year":"2008","journal-title":"Cancer Res."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"1884","DOI":"10.4049\/jimmunol.0900798","article-title":"Novel MHC class I structures on exosomes","volume":"183","author":"Lynch","year":"2009","journal-title":"J. Immunol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1111\/j.1365-2567.2011.03518.x","article-title":"MHC class I dimer formation by alteration of the cellular redox environment and induction of apoptosis","volume":"135","author":"Makhadiyeva","year":"2012","journal-title":"Immunology"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1111\/j.1365-2567.2007.02623.x","article-title":"Synthesis of beta(2)-microglobulin-free, disulphide-linked HLA-G5 homodimers in human placental villous cytotrophoblast cells","volume":"122","author":"Morales","year":"2007","journal-title":"Immunology"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1083\/jcb.98.2.725","article-title":"Interaction between major histocompatibility complex antigens and epidermal growth factor receptors on human cells","volume":"98","author":"Schreiber","year":"1984","journal-title":"J. Cell Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1073\/pnas.84.5.1351","article-title":"Insulin binding to human B lymphoblasts is a function of HLA haplotype","volume":"84","author":"Kittur","year":"1987","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"2463","DOI":"10.1091\/mbc.8.12.2463","article-title":"Interaction of class I human leukocyte antigen (HLA-I) molecules with insulin receptors and its effect on the insulin-signaling cascade","volume":"8","author":"Ramalingam","year":"1997","journal-title":"Mol. Biol. Cell"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/BF00430914","article-title":"Rood Interaction between des-Tyr1-gamma-endorphin and HLA class I molecules: Serological detection of an HLA-A2 subtype","volume":"22","author":"Claas","year":"1985","journal-title":"Immunogenetics"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"3594","DOI":"10.1073\/pnas.88.9.3594","article-title":"Surface appearance and instability of empty H-2 class I molecules under physiological conditions","volume":"88","year":"1991","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"1285","DOI":"10.1002\/eji.1830240607","article-title":"Exogenous peptide ligand influences the expression and half-life of free HLA class I heavy chains ubiquitously detected at the cell surface","volume":"24","author":"Carreno","year":"1994","journal-title":"Eur. J. Immunol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1146\/annurev.immunol.16.1.359","article-title":"NK cell receptors","volume":"16","author":"Lanier","year":"1998","journal-title":"Annu. Rev. Immunol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/978-1-4615-5355-7_2","article-title":"Activating and inhibitory NK cell receptors","volume":"452","author":"Lanier","year":"1998","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1016\/S0092-8674(00)81398-7","article-title":"Follow the leader: NK cell receptors for classical and nonclassical MHC class I","volume":"92","author":"Lanier","year":"1998","journal-title":"Cell"},{"key":"ref_130","doi-asserted-by":"crossref","unstructured":"Debska-Zielkowska, J., Moszkowska, G., Zielinski, M., Zielinska, H., Dukat-Mazurek, A., Trzonkowski, P., and Stefanska, K. (2021). KIR Receptors as Key Regulators of NK Cells Activity in Health and Disease. Cells, 10.","DOI":"10.3390\/cells10071777"},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"3725","DOI":"10.4049\/jimmunol.173.6.3725","article-title":"Stimulatory killer Ig-like receptors modulate T cell activation through DAP12-dependent and DAP12-independent mechanisms","volume":"173","author":"Snyder","year":"2004","journal-title":"J. Immunol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.it.2003.10.011","article-title":"The double life of NK receptors: Stimulation or co-stimulation?","volume":"25","author":"Snyder","year":"2004","journal-title":"Trends Immunol."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1111\/j.1600-065X.1997.tb00945.x","article-title":"Specificity and function of immunoglobulin superfamily NK cell inhibitory and stimulatory receptors","volume":"155","author":"Colonna","year":"1997","journal-title":"Immunol. Rev."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/S0923-2494(97)84220-4","article-title":"Immunoglobulin superfamily inhibitory receptors: From natural killer cells to antigen-presenting cells","volume":"148","author":"Colonna","year":"1997","journal-title":"Res. Immunol."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/1074-7613(95)90025-X","article-title":"Molecular clones of the p58 NK cell receptor reveal immunoglobulin-related molecules with diversity in both the extra- and intracellular domains","volume":"2","author":"Wagtmann","year":"1995","journal-title":"Immunity"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"6313","DOI":"10.1073\/pnas.94.12.6313","article-title":"Peptides isolated from HLA-Cw*0304 confer different degrees of protection from natural killer cell-mediated lysis","volume":"94","author":"Zappacosta","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1034\/j.1600-065X.2001.1810105.x","article-title":"Structure of killer cell immunoglobulin-like receptors and their recognition of the class I MHC molecules","volume":"181","author":"Boyington","year":"2001","journal-title":"Immunol. Rev."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"2972","DOI":"10.1002\/art.10605","article-title":"Cell-surface expression and immune receptor recognition of HLA-B27 homodimers","volume":"46","author":"Kollnberger","year":"2002","journal-title":"Arthritis Rheum."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"1313","DOI":"10.1002\/eji.200635997","article-title":"Interaction of HLA-B27 homodimers with KIR3DL1 and KIR3DL2, unlike HLA-B27 heterotrimers, is independent of the sequence of bound peptide","volume":"37","author":"Kollnberger","year":"2007","journal-title":"Eur. J. Immunol."},{"key":"ref_140","doi-asserted-by":"crossref","unstructured":"Shaw, J., and Kollnberger, S. (2012). New perspectives on the ligands and function of the killer cell immunoglobulin-like receptor KIR3DL2 in health and disease. Front. Immunol., 3.","DOI":"10.3389\/fimmu.2012.00339"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"1699","DOI":"10.4049\/jimmunol.173.3.1699","article-title":"HLA-B27 heavy chain homodimers are expressed in HLA-B27 transgenic rodent models of spondyloarthritis and are ligands for paired Ig-like receptors","volume":"173","author":"Kollnberger","year":"2004","journal-title":"J. Immunol."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/978-1-4419-0298-6_21","article-title":"The role of B27 heavy chain dimer immune receptor interactions in spondyloarthritis","volume":"649","author":"Kollnberger","year":"2009","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"6184","DOI":"10.4049\/jimmunol.1102711","article-title":"HLA-B27 homodimers and free H chains are stronger ligands for leukocyte Ig-like receptor B2 than classical HLA class I","volume":"188","author":"Giles","year":"2012","journal-title":"J. Immunol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"3216","DOI":"10.4049\/jimmunol.1202926","article-title":"KIR3DL2 binds to HLA-B27 dimers and free H chains more strongly than other HLA class I and promotes the expansion of T cells in ankylosing spondylitis","volume":"190","author":"Ridley","year":"2013","journal-title":"J. Immunol."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.molimm.2013.05.243","article-title":"The biochemistry and immunology of non-canonical forms of HLA-B27","volume":"57","author":"Shaw","year":"2014","journal-title":"Mol. Immunol."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"3311","DOI":"10.1182\/blood-2011-03-341065","article-title":"T(H)1, T(H)2, and T(H)17 cells instruct monocytes to differentiate into specialized dendritic cell subsets","volume":"118","author":"Alonso","year":"2011","journal-title":"Blood"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"13028","DOI":"10.1073\/pnas.1000546107","article-title":"Natural killer cells trigger osteoclastogenesis and bone destruction in arthritis","volume":"107","author":"Stein","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1111\/j.0001-2815.2004.00290.x","article-title":"The LILR family: Modulators of innate and adaptive immune pathways in health and disease","volume":"64","author":"Brown","year":"2004","journal-title":"Tissue Antigens"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"656","DOI":"10.1038\/nature11095","article-title":"Inhibitory receptors bind ANGPTLs and support blood stem cells and leukaemia development","volume":"485","author":"Zheng","year":"2012","journal-title":"Nature"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"5192","DOI":"10.4049\/jimmunol.159.11.5192","article-title":"A family of human lymphoid and myeloid Ig-like receptors, some of which bind to MHC class I molecules","volume":"159","author":"Borges","year":"1997","journal-title":"J. Immunol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1189\/jlb.0507314","article-title":"Differential expression of leukocyte immunoglobulin-like receptors on cord-blood-derived human mast cell progenitors and mature mast cells","volume":"83","author":"Tedla","year":"2008","journal-title":"J. Leukoc. Biol."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"2990","DOI":"10.4049\/jimmunol.1003078","article-title":"HLA class I allelic sequence and conformation regulate leukocyte Ig-like receptor binding","volume":"186","author":"Jones","year":"2011","journal-title":"J. Immunol."},{"key":"ref_153","doi-asserted-by":"crossref","unstructured":"Zhang, Z., Hatano, H., Shaw, J., Nordkamp, O.M., Jiang, G., Li, D., and Kollnberger, S. (2015). The Leukocyte Immunoglobulin-Like Receptor Family Member LILRB5 Binds to HLA-Class I Heavy Chains. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0129063"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"308","DOI":"10.4049\/jimmunol.2001165","article-title":"The Structure of a Peptide-Loaded Shark MHC Class I Molecule Reveals Features of the Binding between \u03b22-Microglobulin and H Chain Conserved in Evolution","volume":"207","author":"Wu","year":"2021","journal-title":"J. Immunol."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"742","DOI":"10.1038\/344742a0","article-title":"Beta 2-microglobulin deficient mice lack CD4\u22128+ cytolytic T cells","volume":"344","author":"Zijlstra","year":"1990","journal-title":"Nature"},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1084\/jem.176.3.829","article-title":"Functionally conformed free class I heavy chains exist on the surface of beta 2 microglobulin negative cells","volume":"176","author":"Bix","year":"1992","journal-title":"J. Exp. Med."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"963","DOI":"10.12688\/f1000research.15386.2","article-title":"Major Histocompatibility Complex (MHC) Fragment Numbers Alone\u2014In Atlantic Cod and in General\u2014Do Not Represent Functional Variability","volume":"7","author":"Dijkstra","year":"2018","journal-title":"F1000Res"},{"key":"ref_158","doi-asserted-by":"crossref","unstructured":"Dijkstra, J.M., Grimholt, U., Leong, J., Koop, B.F., and Hashimoto, K. (2013). Comprehensive Analysis of MHC Class II Genes in Teleost Fish Genomes Reveals Dispensability of the Peptide-Loading DM System in a Large Part of Vertebrates. BMC Evol. Biol., 13.","DOI":"10.1186\/1471-2148-13-260"},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Grimholt, U., Tsukamoto, K., Azuma, T., Leong, J., Koop, B.F., and Dijkstra, J.M. (2015). A Comprehensive Analysis of Teleost MHC Class I Sequences. BMC Evol. Biol., 15.","DOI":"10.1186\/s12862-015-0309-1"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1007\/s00251-017-1050-2","article-title":"Conservation of Sequence Motifs Suggests That the Nonclassical MHC Class I Lineages CD1\/PROCR and UT Were Established Before the Emergence of Tetrapod Species","volume":"70","author":"Dijkstra","year":"2018","journal-title":"Immunogenetics"},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1007\/s00251-018-1090-2","article-title":"Ancient Features of the MHC Class II Presentation Pathway, and a Model for the Possible Origin of MHC Molecules","volume":"71","author":"Dijkstra","year":"2019","journal-title":"Immunogenetics"},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0092-8674(84)90068-0","article-title":"The Class II Molecules of the Human and Murine Major Histocompatibility Complex","volume":"36","author":"Kaufman","year":"1984","journal-title":"Cell"},{"key":"ref_163","first-page":"131","article-title":"Vertebrates and the Evolution of the Major Histocompatibility Complex (MHC) Class I and Class II Molecules","volume":"81","author":"Kaufman","year":"1988","journal-title":"Verh. Dtsch. Zool. Ges."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"1532","DOI":"10.4049\/jimmunol.148.5.1532","article-title":"Different Features of the MHC Class I Heterodimer Have Evolved at Different -Rates. Chicken B-F and Beta 2-Microglobulin Sequences Reveal Invariant Surface Residues","volume":"148","author":"Kaufman","year":"1992","journal-title":"J. Immunol."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"2209","DOI":"10.1073\/pnas.89.6.2209","article-title":"Identification of a shark sequence resembling the major histocompatibility complex class I alpha 3 domain","volume":"89","author":"Hashimoto","year":"1992","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"3475926","DOI":"10.1155\/2017\/3475926","article-title":"HLA Class Ia and Ib Polyreactive Anti-HLA-E IgG2a Monoclonal Antibodies (TFL-006 and TFL-007) Suppress Anti-HLA IgG Production by CD19+ B Cells and Proliferation of CD4+ T Cells While Upregulating Tregs","volume":"2017","author":"Ravindranath","year":"2017","journal-title":"J. Immunol. Res."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"18","DOI":"10.3390\/antib11010018","article-title":"Ramifications of the HLA-I Allelic Reactivity of Anti-HLA-E*01:01 and Anti-HLA-E*01:03 Heavy Chain Monoclonal Antibodies in Comparison with Anti-HLA-I IgG Reactivityin Non-Alloimmunized Males, Melanoma-Vaccine Recipients, and End-Stage Renal Disease Patients","volume":"11","author":"Ravindranath","year":"2022","journal-title":"Antibodies"}],"container-title":["Current Issues in Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1467-3045\/46\/7\/416\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T15:10:09Z","timestamp":1760109009000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1467-3045\/46\/7\/416"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,7,4]]},"references-count":167,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2024,7]]}},"alternative-id":["cimb46070416"],"URL":"https:\/\/doi.org\/10.3390\/cimb46070416","relation":{},"ISSN":["1467-3045"],"issn-type":[{"value":"1467-3045","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,7,4]]}}}