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CD80
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<h2 id="structure">Structure</h2> <p>CD80 is a member of the <a href="/facts/B7_(protein)/UyqjJrGB">B7</a> family, which consists of molecules present at <a href="/facts/Antigen-presenting_cell/No0wr2uU">APCs</a> and their receptors present on the <a href="/facts/T_cell/5z6PQ9UN">T-cells</a>.<a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a> CD80 is present specifically on <a href="/facts/Dendritic_cell/19KBuKFK">DC</a>, activated <a href="/facts/B_cell/6Y1tgwwp">B-cells</a>, and <a href="/facts/Macrophage/z7hkQu3o">macrophages</a>, but also <a href="/facts/T_cell/5z6PQ9UN">T-cells</a>.<a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a><a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> CD80 is also a <a href="/facts/Transmembrane_protein/0vQH5HrM">transmembrane</a> <a href="/facts/Glycoprotein/bKPzzW7m">glycoprotein</a> and a member of the <a href="/facts/Immunoglobulin_superfamily/Wveu4BIL">Ig superfamily</a>.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> It is composed of 288 <a href="/facts/Amino_acid/WDxYwBny">amino acids</a>, and its mass is 33 <a href="/facts/Dalton_(unit)/IMATU3vf">kDa</a>.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> It consists of two Ig-like extracellular domains (208 AA), a transmembrane helical segment (21 AA), and a short cytoplasmic tail (25 AA).<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a><a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a><a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> The Ig-like extracellular domains are formed by single V-type and C2-type domains.<a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a><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> It is expressed as both <a href="/facts/Monomer/Gw3CltMb">monomers</a> or <a href="/facts/Protein_dimer/c8LwFJPG">dimers</a>, but predominantly dimers.<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> These two forms exist in dynamic equilibrium.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> </p><p>CD80 shares 25% of sequences with <a href="/facts/CD86/VUSlNjoW">CD86</a>; however, CD80 has a ten-fold higher <a href="/facts/Chemical_affinity/IEkqFIP4">affinity</a> for <a href="/facts/CD28/jutfUEHw">CD28</a> and <a href="/facts/CTLA-4/7nhf4Nq6">CTLA-4</a> than <a href="/facts/CD86/VUSlNjoW">CD86</a>. Moreover, CD80 interacts with its ligand with faster binding kinetics and slower dissociation constants than <a href="/facts/CD86/VUSlNjoW">CD86</a>. Both human CD80 and CD86 are located at <a href="/facts/Chromosome_3/2lHrRbHd">chromosome 3</a>; the exact region is 3q13.3-q21.<a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a> </p><p><a href="/facts/Human/rlcTENsr">Human</a> and <a href="/facts/Murinae/7ds8prdo">murine</a> CD80 share approximately 44% of sequences. Also both human and murine CD80 are able to cross-react with both human and murine CD28. This indicates that the binding site of CD80 is conserved.<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a><a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> </p> <h2 id="function">Function</h2> <p>CD80 can be found on the surface of various <a href="/facts/Immune_cells/vsDunUK0">immune cells</a>, including <a href="/facts/B_cells/6Y1tgwwp">B-cells</a>, <a href="/facts/Monocytes/FW4NNMKM">monocytes</a>, or T-cells, but most typically at <a href="/facts/Antigen-presenting_cells/No0wr2uU">antigen-presenting cells</a> (APCs) such as <a href="/facts/Dendritic_cells/19KBuKFK">dendritic cells</a>.<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a><a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a><a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a> CD80 has a crucial role in modulating T-cell immune function as a checkpoint protein at the <a href="/facts/Immunological_synapse/3e8dcntv">immunological synapse</a>.<a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a> </p><p>CD80 is the ligand for the proteins <a href="/facts/CD28/jutfUEHw">CD28</a> (for autoregulation and intercellular association) and <a href="/facts/CTLA-4/7nhf4Nq6">CTLA-4</a> (for attenuation of regulation and cellular disassociation) found on the surface of <a href="/facts/T-cells/5z6PQ9UN">T-cells</a>.<a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a><a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a> Interaction of CD80 with CD28 triggers <a href="/facts/Co-stimulation/1s3weomD">costimulatory</a> signals and results in enhanced and sustained T-cell activation. In contrast, contrary interaction of CD80 with CTLA-4 inhibits parts of T-cell effector function. These two <a href="/facts/Ligand/6etB3qeW">ligands</a> are structurally homologous, and they compete with each other for <a href="/facts/Binding_site/gd9r9Rw7">binding sites</a>.<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a> However, the bond with CTLA-4 has up to 2500 fold higher <a href="/facts/Avidity/zrA1tXeI">avidity</a> than with CD28.<a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a> This illustrates that inhibitory interaction with CTLA-4 is predominant.<a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a> </p><p>CD80 binds to <a href="/facts/CD28/jutfUEHw">CD28</a> and <a href="/facts/CTLA-4/7nhf4Nq6">CTLA-4</a> with lower <a href="/facts/Chemical_affinity/IEkqFIP4">affinity</a> and fast binding kinetics (Kd = 4 μM for CD28 and 0.42 μM for CTLA-4), allowing for quick interactions between the communicating cells.<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a> These interactions result in an important costimulatory signal in the <a href="/facts/Immunological_synapse/3e8dcntv">immunological synapse</a> between <a href="/facts/Antigen-presenting_cells/No0wr2uU">antigen-presenting cells</a>, <a href="/facts/B-cells/6Y1tgwwp">B-cells</a>, <a href="/facts/Dendritic_cells/19KBuKFK">dendritic cells</a> and <a href="/facts/T-cells/5z6PQ9UN">T-cells</a> that result in T and B-cell activation, proliferation and differentiation.<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a> </p><p>When stimulated by CD80, <a href="/facts/T_helper_cell/nhwAYkDj">T helper cells</a> preferentially differentiate into <a href="/facts/T_helper_cell/nhwAYkDj">Th1 cells</a>.<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> CD80 is an essential component in <a href="/facts/Dendritic_cell/19KBuKFK">dendritic cell</a> licensing and <a href="/facts/Cytotoxic_T-cell/e5eRg3XJ">cytotoxic T-cell</a> activation. When the major histocompatibility complex class II (<a href="/facts/MHC_class_II/nQ4hkBgW">MHC class II</a>)-peptide complex on a <a href="/facts/Dendritic_cell/19KBuKFK">dendritic cell</a> interacts with the receptor on a <a href="/facts/T_helper_cell/nhwAYkDj">T helper cell</a>, CD80 is up-regulated, licensing the <a href="/facts/Dendritic_cell/19KBuKFK">dendritic cell</a> and allowing for interaction between the <a href="/facts/Dendritic_cell/19KBuKFK">dendritic cell</a> and <a href="/facts/Cytotoxic_T_cell/e5eRg3XJ">CD 8+ T-cells</a> via <a href="/facts/CD28/jutfUEHw">CD28</a>. This helps to signal the T-cell differentiation into a <a href="/facts/Cytotoxic_T-cell/e5eRg3XJ">cytotoxic T-cell</a>.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a><a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a> The expression of CD80, as well as <a href="/facts/CD86/VUSlNjoW">CD86</a>, is increased by the presence of <a href="/facts/Microorganism/pysJYcE2">microbes</a> and <a href="/facts/Cytokine/SdWpWABH">cytokines</a>, which is the consequence of the presence of microbes. This mechanism ensures that costimulatory molecules for <a href="/facts/T_cell/5z6PQ9UN">T-cells</a> are present at the right time.<a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> </p><p>CD80, often in tandem with <a href="/facts/CD86/VUSlNjoW">CD86</a>, plays a large and diverse role in regulating both the <a href="/facts/Adaptive_immune_system/bc7zT1Jo">adaptive</a> and the <a href="/facts/Innate_immune_system/lz4BpSQI">innate immune system</a>. As mentioned above, this protein is crucial for immune cell activation in response to <a href="/facts/Pathogen/axMXtV1M">pathogens</a>. The interaction of CD80 with CD28, together with <a href="/facts/T-cell_receptor/zDqyWyG4">TCR</a> and <a href="/facts/Major_histocompatibility_complex/UEPsjx3u">MHC</a> interaction, results in activation of nuclear factor-κB (<a href="/facts/NF-%CE%BAB/qcpt5vuQ">NF-ⲕB</a>), mitogen-activated protein kinase (<a href="/facts/Mitogen-activated_protein_kinase/YdDS5KkU">MAPK</a>), and the calcium-calcineurin pathway. These changes initiate the production of numerous factors, <a href="/facts/Cytokine/SdWpWABH">cytokines</a>, and <a href="/facts/Chemokine/YrySj9eD">chemokines</a> by T-cells. Noteworthy is the production of interleukin 2 (<a href="/facts/Interleukin_2/hH1DXu2W">IL-2</a>) as well as ɑ-chain of <a href="/facts/IL2RA/Woa9fxbk">CD25</a> (which is a receptor of IL-2), <a href="/facts/CD154/Tx7z1XPp">CD40 ligand</a>, tumor necrosis factor-α (<a href="/facts/Tumor_necrosis_factor/7NWNIaCb">TNFα</a>), TNF-β, and interferon-γ (<a href="/facts/Interferon_gamma/r51CXVph">IFN-γ</a>). T-cells also increase the production of <a href="/facts/Macrophage_inflammatory_protein/RPAp5XY2">macrophage inflammatory proteins</a> 1α and 1β (MIP-α1 and MIP-1β) and prevent <a href="/facts/Apoptosis/8dVzSm4y">apoptosis</a> by induction of anti-apoptotic protein expression (e.g., <a href="/facts/Bcl-x_interacting_domain/ClugpM3M">Bcl-X</a> and <a href="/facts/Bcl-2/I34R1guW">Bcl-2</a>).<a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a><a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a><a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a><a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a><a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a> CD80 interaction with <a href="/facts/CD28/jutfUEHw">CD28</a> also further stimulates <a href="/facts/Dendritic_cells/19KBuKFK">dendritic cells</a>, enhancing <a href="/facts/Cytokine/SdWpWABH">cytokine</a> production, specifically <a href="/facts/Interleukin_6/FVQeAWkt">IL-6</a>, a pro-inflammatory molecule.<a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a><a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> <a href="/facts/Neutrophils/KL6oTt8b">Neutrophils</a> can also activate <a href="/facts/Macrophages/z7hkQu3o">macrophages</a> with CD80 via <a href="/facts/CD28/jutfUEHw">CD28</a>.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> Last but not least, the interaction of CD80 and CD28 enhances cell-cycle progression by upregulating the expression levels of <a href="/facts/Cyclin_D/XrAPyeCw">D-cyclin</a>.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> </p><p>In contrast to the stimulatory interaction with <a href="/facts/CD28/jutfUEHw">CD28</a>, CD80 also regulates the immune system through an inhibitory interaction with <a href="/facts/CTLA-4/7nhf4Nq6">CTLA-4</a>. <a href="/facts/Dendritic_cells/19KBuKFK">Dendritic cells</a> are suppressed by a CTLA-4-CD80 interaction,<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> and this interaction also promotes the suppressive effects of <a href="/facts/Regulatory_T_cell/1ECRAGMg">regulatory T cells</a>, which can prevent an immune response to <a href="/facts/Self-antigen/hvDqPHTP">self-antigen</a>.<a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> </p><p>In addition to interactions with <a href="/facts/CD28/jutfUEHw">CD28</a> and <a href="/facts/CTLA-4/7nhf4Nq6">CTLA-4</a>, CD80 is also thought to interact with a separate ligand on <a href="/facts/Natural_Killer_cell/A3THWxrJ">Natural Killer cells</a>, triggering the Natural Killer cell-mediated cell death of the CD80 carrier.<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> CD80 may also play a role in the negative regulation of effector and memory T-cells. If the interaction between an <a href="/facts/Antigen-presenting_cell/No0wr2uU">antigen-presenting cell</a> and a <a href="/facts/T-cell/5z6PQ9UN">T-cell</a> is stable enough, the T-cell can remove the CD80 from the antigen-presenting cell by a mechanism dubbed <a href="/facts/Trans-endocytosis/673f9qXv">trans-endocytosis</a>. Under the right conditions, this transfer of the CD80 may induce T-cell <a href="/facts/Apoptosis/8dVzSm4y">apoptosis</a>.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> Finally, CD80 signaling on activated <a href="/facts/B-cells/6Y1tgwwp">B-cells</a> may regulate antibody secretion during <a href="/facts/Infection/182XrncP">infection</a>.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> </p><p>Another ligand of CD80 is programmed death-ligand 1 (<a href="/facts/PD-L1/0jyfJ2nE">PD-L1</a>), expressed on the surface of T-cells, B-cells, DCs, and macrophages. This interaction is inhibiting and causes a reduction in T-cell activation as well as reduction of cytokine production. Its dissociation constant with CD80 is between the CD28 and CTLA-40 (Kd = 1.4 μM).<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a><a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> </p> <h2 id="clinical-significance">Clinical significance</h2> <p>The complicated role CD80 plays in immune system regulation presents an opportunity for CD80 interactions to go rogue in various diseases. The up-regulation of CD80 has been linked to various <a href="/facts/Autoimmune_diseases/sojBeQ7t">autoimmune diseases</a>, including <a href="/facts/Multiple_sclerosis/vghXl0IQ">multiple sclerosis</a>,<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> <a href="/facts/Systemic_lupus_erythematosus/kfHYawre">systemic lupus erythematosus</a><a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> and <a href="/facts/Sepsis/n52bkffJ">sepsis</a><a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a> (which may partly be due to over-active T-cells), and CD80 has also been shown to help spread of <a href="/facts/HIV/ulY2rgzu">HIV</a> infection in the body.<a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> CD80 is also linked to various <a href="/facts/Cancer/PVW6n1D0">cancers</a>, though some experience CD80 induced tolerance via possible <a href="/facts/Regulatory_T_cell/1ECRAGMg">regulatory T-cell</a> interaction.<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> Others experience inhibited growth and <a href="/facts/Metastasis/4dadbm6z">metastasis</a>-related to CD80 up-regulation,<a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a> further exemplifies the complicated role CD80 plays. </p><p>The triggering of <a href="/facts/Natural_Killer_cell/A3THWxrJ">Natural Killer cell</a>-mediated death via CD80 interactions has been explored as possible cancer <a href="/facts/Immunotherapy/hgq3KQaJ">immunotherapy</a> by inducing CD80 expression on tumor cells.<a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></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/CD86/VUSlNjoW">CD86</a></li> <li><a href="/facts/CD28/jutfUEHw">CD28</a></li></ul> <h2 id="external-links">External links</h2> <ul><li>Human <a href="https://genome.ucsc.edu/cgi-bin/hgTracks?db=hg38&singleSearch=knownCanonical&position=CD80"><i>CD80</i></a> genome location and <a href="https://genome.ucsc.edu/cgi-bin/hgGene?db=hg38&hgg_type=knownGene&hgg_gene=CD80"><i>CD80</i></a> gene details page in the <a href="/facts/UCSC_Genome_Browser/kwumPyLb">UCSC Genome Browser</a>.</li> <li>Overview of all the structural information available in the <a href="/facts/Protein_Data_Bank/oUhq4ABD">PDB</a> for <a href="/facts/UniProt/lfDDHjAl">UniProt</a>: <i><a href="https://www.ebi.ac.uk/pdbe/pdbe-kb/proteins/P33681">P33681</a></i> (T-lymphocyte activation antigen CD80) at the <a href="/facts/PDBe-KB/p1WX7lPH">PDBe-KB</a>.</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>McKusick, V. A., & Converse, P. J. (2016, August 05). CD80 Antigen; CD80. Retrieved May 29, 2019 <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Peach RJ, Bajorath J, Naemura J, Leytze G, Greene J, Aruffo A, Linsley PS (September 1995). "Both extracellular immunoglobin-like domains of CD80 contain residues critical for binding T cell surface receptors CTLA-4 and CD28". The Journal of Biological Chemistry. 270 (36): 21181–7. doi:10.1074/jbc.270.36.21181. PMID 7545666. <a href="https://doi.org/10.1074%2Fjbc.270.36.21181" target="_blank">https://doi.org/10.1074%2Fjbc.270.36.21181</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>"CD80 - T-lymphocyte activation antigen CD80 precursor - Homo sapiens (Human) - CD80 gene & protein". www.uniprot.org. Retrieved 2021-06-09. <a href="https://www.uniprot.org/uniprot/P33681" target="_blank">https://www.uniprot.org/uniprot/P33681</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>"CD80 - T-lymphocyte activation antigen CD80 precursor - Homo sapiens (Human) - CD80 gene & protein". www.uniprot.org. Retrieved 2021-06-09. <a href="https://www.uniprot.org/uniprot/P33681" target="_blank">https://www.uniprot.org/uniprot/P33681</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>"CD80 - T-lymphocyte activation antigen CD80 precursor - Homo sapiens (Human) - CD80 gene & protein". www.uniprot.org. Retrieved 2021-06-09. <a href="https://www.uniprot.org/uniprot/P33681" target="_blank">https://www.uniprot.org/uniprot/P33681</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Abbas AK (2021). Cellular and molecular immunology. Elsevier. ISBN 978-0-323-75749-2. OCLC 1173994133. <a href="978-0-323-75749-2" target="_blank">978-0-323-75749-2</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. ISBN 978-0-12-802585-7. <a href="978-0-12-802585-7" target="_blank">978-0-12-802585-7</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Peach RJ, Bajorath J, Naemura J, Leytze G, Greene J, Aruffo A, Linsley PS (September 1995). "Both extracellular immunoglobin-like domains of CD80 contain residues critical for binding T cell surface receptors CTLA-4 and CD28". The Journal of Biological Chemistry. 270 (36): 21181–7. doi:10.1074/jbc.270.36.21181. PMID 7545666. <a href="https://doi.org/10.1074%2Fjbc.270.36.21181" target="_blank">https://doi.org/10.1074%2Fjbc.270.36.21181</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Bhatia S, Edidin M, Almo SC, Nathenson SG (April 2006). "B7-1 and B7-2: similar costimulatory ligands with different biochemical, oligomeric and signaling properties". Immunology Letters. 104 (1–2): 70–5. doi:10.1016/j.imlet.2005.11.019. PMID 16413062. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>Mir MA (2015). "Introduction to Costimulation and Costimulatory Molecules". Developing Costimulatory Molecules for Immunotherapy of Diseases. Elsevier: 1–43. doi:10.1016/b978-0-12-802585-7.00001-7. 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