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CD74
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<h2 id="function">Function</h2> <p>The nascent <a href="/facts/Major_histocompatibility_complex/UEPsjx3u">MHC</a> class II protein in the <a href="/facts/Endoplasmic_reticulum/DVbLA7GE">rough endoplasmic reticulum</a> (RER) binds a segment of the invariant chain (Ii; a trimer) in order to shape the peptide-binding groove and prevent the formation of a closed conformation. </p><p>The invariant chain also facilitates the export of MHC class II from the RER in a vesicle. The signal for endosomal targeting resides in the cytoplasmic tail of the invariant chain. This fuses with a late <a href="/facts/Endosome/CYsyuJ75">endosome</a> containing the endocytosed antigen proteins (from the exogenous pathway). Binding to Ii ensures that no antigen peptides from the endogenous pathway meant for <a href="/facts/MHC_class_I/5VsuyBc4">MHC class I</a> molecules accidentally bind to the groove of MHC class II molecules.<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> The Ii is then cleaved by <a href="/facts/Cathepsin_S/ceuTfrBC">cathepsin S</a> (<a href="/facts/Cathepsin_L1/RIftLTSH">cathepsin L</a> in <a href="/facts/Cortical_thymic_epithelial_cells/R97wotWs">cortical thymic epithelial cells</a>), leaving only a small fragment called <a href="/facts/CLIP_(protein)/FfRXP0sg">CLIP</a> remaining bound to the groove of MHC class II molecules. The rest of the Ii is degraded.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> CLIP blocks peptide-binding until <a href="/facts/HLA-DM/bp8WRWW6">HLA-DM</a> interacts with MHC II, releasing CLIP and allowing other peptides to bind. In some cases, CLIP dissociates without any further molecular interactions, but in other cases the binding to the MHC is more stable.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> </p><p>The stable MHC class II + antigen complex is then <a href="/facts/Antigen_presentation/RMEAcYwR">presented</a> on the cell surface. Without CLIP, MHC class II aggregates disassemble and/or denature in the endosomes, and proper antigen presentation is impaired.<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> </p> <h2 id="clinical-significance">Clinical significance</h2> <h3>Vaccine adjuvant</h3> <p>The Ii molecule—fused with a <a href="/facts/Viral_vector/sqkIdcqo">viral vector</a> to a <a href="/facts/Conserved_sequence/YAxy2Xa2">conserved region</a> of the <a href="/facts/Hepatitis_C_virus/DFUwAzlB">Hepatitis C virus</a> (HCV) genome—has been tested as an <a href="/facts/Adjuvant/jnRqeBnG">adjuvant</a> for a <a href="/facts/Hepatitis_C_vaccine/wzXnLIqa">HCV vaccine</a> in a cohort of 17 healthy human volunteers. This experimental vaccine was well-tolerated, and those who received the adjuvanted vaccine had stronger anti-HCV immune responses (enhanced magnitude, breadth and proliferative capacity of anti-HCV-specific T-cells) compared with volunteers who received the vaccine that lacked the Ii adjuvant.<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> </p><p>The Ii molecule might also prove to be useful as an adjuvant for a future <a href="/facts/COVID-19_vaccine/obToxWTT">vaccine for the SARS-CoV-2 virus</a>, if this enhancing effect can be demonstrated to apply to the appropriate antigen(s).<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> </p> <h3>Cancer</h3> <p>Found on a number of cancer cell types. Possible cancer therapy target. See <a href="/facts/Milatuzumab/oGW2WkBN">milatuzumab</a>. </p> <h3>Axial spondyloarthritis</h3> <p><a href="/facts/Autoantibodies/gkMT0XyQ">Autoantibodies</a> against CD74 have been identified as promising <a href="/facts/Biomarkers/nk2LgXtq">biomarkers</a> in the early diagnosis of the <a href="/facts/Autoimmune_disease/sojBeQ7t">autoimmune disease</a> called <a href="/facts/Axial_spondyloarthritis/1875YJwt">axial spondyloarthritis</a> (<a href="/facts/Non-radiographic_axial_spondyloarthritis/1875YJwt">non-radiographic axial spondyloarthritis</a> and <a href="/facts/Radiographic_axial_spondyloarthritis/1875YJwt">radiographic axial spondyloarthritis</a> / <a href="/facts/Ankylosing_spondylitis/tHgAlJNd">Ankylosing spondylitis</a>). <a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> </p> <h2 id="interactions">Interactions</h2> <p>CD74 receptor <a href="/facts/Protein-protein_interaction/etvm9Wmg">interacts</a> with the cytokine <a href="/facts/Macrophage_migration_inhibitory_factor/QIBya53j">Macrophage migration inhibitory factor</a> (MIF) to mediate some of its functions.<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a><a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a><a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a><a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a><a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a><a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> </p> <h2 id="recovery-functions">Recovery functions</h2> <p>CD74 receptor is expressed on the surface of different cell types. Interaction between MIF cytokine and its cell membrane receptor CD74 activates pro-survival and proliferative pathways that protect against injury and promote healing in different parts of the body.<a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a> </p> <h2 id="history">History</h2> <p>The invariant chain was first described by Patricia P. Jones, Donal B. Murphy, Derek Hewgill, and <a href="/facts/Hugh_McDevitt/8qvphyuY">Hugh McDevitt</a> at Stanford.<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> The nomenclature "Ii" comes from an Ix-based naming system (<i>I</i> for <i>I</i>mmune) that predates the naming of the <a href="/facts/Major_histocompatibility_complex/UEPsjx3u">Major Histocompatibility Complex</a>. </p> <h2 id="see-also">See also</h2> <ul><li><a href="/facts/Cluster_of_differentiation/cLEuDQId">Cluster of differentiation</a></li> <li><a href="/facts/Milatuzumab/oGW2WkBN">Milatuzumab</a> the first Mab to target CD74</li></ul> <h2 id="further-reading">Further reading</h2> <ul><li>Stove V, Verhasselt B (January 2006). "Modelling thymic HIV-1 Nef effects". <i>Current HIV Research</i>. 4 (1): 57–64. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.2174%2F157016206775197583">10.2174/157016206775197583</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/16454711">16454711</a>.</li> <li>Riberdy JM, Newcomb JR, Surman MJ, Barbosa JA, Cresswell P (December 1992). "HLA-DR molecules from an antigen-processing mutant cell line are associated with invariant chain peptides". <i>Nature</i>. 360 (6403): 474–477. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1992Natur.360..474R">1992Natur.360..474R</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2F360474a0">10.1038/360474a0</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/1448172">1448172</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:4338656">4338656</a>.</li> <li>Bakke O, Dobberstein B (November 1990). <a href="http://archiv.ub.uni-heidelberg.de/volltextserver/9193/1/27.MHC_class_II_associated_invariant_chain.pdf">"MHC class II-associated invariant chain contains a sorting signal for endosomal compartments"</a> (PDF). <i>Cell</i>. 63 (4): 707–716. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2F0092-8674%2890%2990137-4">10.1016/0092-8674(90)90137-4</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/2121367">2121367</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:19993336">19993336</a>.</li> <li>Marks MS, Blum JS, Cresswell P (September 1990). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2116304">"Invariant chain trimers are sequestered in the rough endoplasmic reticulum in the absence of association with HLA class II antigens"</a>. <i>The Journal of Cell Biology</i>. 111 (3): 839–855. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1083%2Fjcb.111.3.839">10.1083/jcb.111.3.839</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2116304">2116304</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/2391366">2391366</a>.</li> <li>Spiro RC, Quaranta V (October 1989). <a href="https://doi.org/10.4049%2Fjimmunol.143.8.2589">"The invariant chain is a phosphorylated subunit of class II molecules"</a>. <i>Journal of Immunology</i>. 143 (8): 2589–2594. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.4049%2Fjimmunol.143.8.2589">10.4049/jimmunol.143.8.2589</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/2507633">2507633</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:6755576">6755576</a>.</li> <li>O'Sullivan DM, Noonan D, Quaranta V (August 1987). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2189580">"Four Ia invariant chain forms derive from a single gene by alternate splicing and alternate initiation of transcription/translation"</a>. <i>The Journal of Experimental Medicine</i>. 166 (2): 444–460. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1084%2Fjem.166.2.444">10.1084/jem.166.2.444</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2189580">2189580</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/3036998">3036998</a>.</li> <li>Genuardi M, Saunders GF (1988). "Localization of the HLA class II-associated invariant chain gene to human chromosome band 5q32". <i>Immunogenetics</i>. 28 (1): 53–56. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1007%2FBF00372530">10.1007/BF00372530</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/3132422">3132422</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:2418453">2418453</a>.</li> <li>O'Sullivan DM, Larhammar D, Wilson MC, Peterson PA, Quaranta V (June 1986). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC323758">"Structure of the human Ia-associated invariant (gamma)-chain gene: identification of 5' sequences shared with major histocompatibility complex class II genes"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. 83 (12): 4484–4488. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1986PNAS...83.4484O">1986PNAS...83.4484O</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1073%2Fpnas.83.12.4484">10.1073/pnas.83.12.4484</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC323758">323758</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/3459184">3459184</a>.</li> <li>Koch N, Hämmerling GJ (October 1985). "Ia-associated invariant chain is fatty acylated before addition of sialic acid". <i>Biochemistry</i>. 24 (22): 6185–6190. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1021%2Fbi00343a023">10.1021/bi00343a023</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/3866610">3866610</a>.</li> <li>Claesson L, Peterson PA (June 1983). "Association of human gamma chain with class II transplantation antigens during intracellular transport". <i>Biochemistry</i>. 22 (13): 3206–3213. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1021%2Fbi00282a026">10.1021/bi00282a026</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/6576808">6576808</a>.</li> <li>Strubin M, Mach B, Long EO (April 1984). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC557440">"The complete sequence of the mRNA for the HLA-DR-associated invariant chain reveals a polypeptide with an unusual transmembrane polarity"</a>. <i>The EMBO Journal</i>. 3 (4): 869–872. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1002%2Fj.1460-2075.1984.tb01898.x">10.1002/j.1460-2075.1984.tb01898.x</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC557440">557440</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/6586420">6586420</a>.</li> <li>Kvist S, Wiman K, Claesson L, Peterson PA, Dobberstein B (May 1982). <a href="http://archiv.ub.uni-heidelberg.de/volltextserver/9169/1/10.Membrane_insertion_and_oligomeric_assembly.pdf">"Membrane insertion and oligomeric assembly of HLA-DR histocompatibility antigens"</a> (PDF). <i>Cell</i>. 29 (1): 61–69. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2F0092-8674%2882%2990090-3">10.1016/0092-8674(82)90090-3</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/6955026">6955026</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:9066996">9066996</a>.</li> <li>Machamer CE, Cresswell P (December 1982). "Biosynthesis and glycosylation of the invariant chain associated with HLA-DR antigens". <i>Journal of Immunology</i>. 129 (6): 2564–2569. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.4049%2Fjimmunol.129.6.2564">10.4049/jimmunol.129.6.2564</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/6982931">6982931</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:42620266">42620266</a>.</li> <li>Ghosh P, Amaya M, Mellins E, Wiley DC (November 1995). "The structure of an intermediate in class II MHC maturation: CLIP bound to HLA-DR3". <i>Nature</i>. 378 (6556): 457–462. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1995Natur.378..457G">1995Natur.378..457G</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2F378457a0">10.1038/378457a0</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/7477400">7477400</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:4275956">4275956</a>.</li> <li>Bijlmakers MJ, Benaroch P, Ploegh HL (August 1994). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2191624">"Mapping functional regions in the lumenal domain of the class II-associated invariant chain"</a>. <i>The Journal of Experimental Medicine</i>. 180 (2): 623–629. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1084%2Fjem.180.2.623">10.1084/jem.180.2.623</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2191624">2191624</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/7519244">7519244</a>.</li> <li>Brown JH, Jardetzky TS, Gorga JC, Stern LJ, Urban RG, Strominger JL, Wiley DC (July 1993). "Three-dimensional structure of the human class II histocompatibility antigen HLA-DR1". <i>Nature</i>. 364 (6432): 33–39. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1993Natur.364...33B">1993Natur.364...33B</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2F364033a0">10.1038/364033a0</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/8316295">8316295</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:4248668">4248668</a>.</li> <li>Naujokas MF, Morin M, Anderson MS, Peterson M, Miller J (July 1993). "The chondroitin sulfate form of invariant chain can enhance stimulation of T cell responses through interaction with CD44". <i>Cell</i>. 74 (2): 257–268. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2F0092-8674%2893%2990417-O">10.1016/0092-8674(93)90417-O</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/8343954">8343954</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:10295196">10295196</a>.</li> <li>Roche PA, Teletski CL, Stang E, Bakke O, Long EO (September 1993). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC47401">"Cell surface HLA-DR-invariant chain complexes are targeted to endosomes by rapid internalization"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. 90 (18): 8581–8585. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1993PNAS...90.8581R">1993PNAS...90.8581R</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1073%2Fpnas.90.18.8581">10.1073/pnas.90.18.8581</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC47401">47401</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/8397411">8397411</a>.</li></ul> <h2 id="external-links">External links</h2> <ul><li><a href="https://meshb.nlm.nih.gov/record/ui?name=CD74+protein%2C+human">CD74+protein,+human</a> at the U.S. National Library of Medicine <a href="/facts/Medical_Subject_Headings/C57g2JuF">Medical Subject Headings</a> (MeSH)</li> <li><a href="http://www.bio.davidson.edu/Courses/immunology/Students/spring2006/McCracken/HLA-DM.html">Overview</a> at <a href="/facts/Davidson_College/FIMeRLFL">Davidson College</a> (student generated)</li> <li><a href="http://www.cryst.bbk.ac.uk/pps97/assignments/projects/coadwell/007.htm">School of Crystallography</a> The Invariant Chain</li> <li>Human <a href="https://genome.ucsc.edu/cgi-bin/hgTracks?db=hg38&singleSearch=knownCanonical&position=CD74"><i>CD74</i></a> genome location and <a href="https://genome.ucsc.edu/cgi-bin/hgGene?db=hg38&hgg_type=knownGene&hgg_gene=CD74"><i>CD74</i></a> gene details page in the <a href="/facts/UCSC_Genome_Browser/kwumPyLb">UCSC Genome Browser</a>.</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Claesson L, Larhammar D, Rask L, Peterson PA (December 1983). 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