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<span>Gene</span> </div> </a> <ul id="toc-Gene-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Function</span> </div> </a> <ul id="toc-Function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Regulation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Regulation</span> </div> </a> <ul id="toc-Regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Clinical significance</span> </div> </a> <button aria-controls="toc-Clinical_significance-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Clinical significance subsection</span> </button> <ul id="toc-Clinical_significance-sublist" class="vector-toc-list"> <li id="toc-Role_in_carcinogenesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_in_carcinogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Role in carcinogenesis</span> </div> </a> <ul id="toc-Role_in_carcinogenesis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Clinical_use" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_use"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Clinical use</span> </div> </a> <button aria-controls="toc-Clinical_use-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Clinical use subsection</span> </button> <ul id="toc-Clinical_use-sublist" class="vector-toc-list"> <li id="toc-Biomarker_for_cancer_types" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Biomarker_for_cancer_types"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Biomarker for cancer types</span> </div> </a> <ul id="toc-Biomarker_for_cancer_types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-p16_FISH" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#p16_FISH"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>p16 FISH</span> </div> </a> <ul id="toc-p16_FISH-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-p16_immunochemistry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#p16_immunochemistry"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>p16 immunochemistry</span> </div> </a> <ul id="toc-p16_immunochemistry-sublist" class="vector-toc-list"> <li id="toc-Gynecologic_cancers" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Gynecologic_cancers"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3.1</span> <span>Gynecologic cancers</span> </div> </a> <ul id="toc-Gynecologic_cancers-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Urinary_bladder_SCCs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Urinary_bladder_SCCs"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span>Urinary bladder SCCs</span> </div> </a> <ul id="toc-Urinary_bladder_SCCs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_cellular_senescence" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_in_cellular_senescence"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.5</span> <span>Role in cellular senescence</span> </div> </a> <ul id="toc-Role_in_cellular_senescence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_neurogenesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_in_neurogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.6</span> <span>Role in neurogenesis</span> </div> </a> <ul id="toc-Role_in_neurogenesis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Discovery" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Discovery"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Discovery</span> </div> </a> <ul id="toc-Discovery-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_carcinogenesis_2" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Role_in_carcinogenesis_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Role in carcinogenesis</span> </div> </a> <ul id="toc-Role_in_carcinogenesis_2-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Interactions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Interactions"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Interactions</span> </div> </a> <ul id="toc-Interactions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" 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class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Mammalian protein found in humans</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">For other uses, see <a href="/wiki/P16_(disambiguation)" class="mw-disambig" title="P16 (disambiguation)">P16 (disambiguation)</a>.</div> <table class="infobox" style="width:26.4em"><tbody><tr><th colspan="4" style="text-align:center;font-size:125%;font-weight:bold">CDKN2A</th></tr><tr><td colspan="4" style="text-align:center"><span typeof="mw:File"><a href="/wiki/File:Protein_CDKN2A_PDB_1a5e.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Protein_CDKN2A_PDB_1a5e.png/250px-Protein_CDKN2A_PDB_1a5e.png" decoding="async" width="250" height="137" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Protein_CDKN2A_PDB_1a5e.png/375px-Protein_CDKN2A_PDB_1a5e.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Protein_CDKN2A_PDB_1a5e.png/500px-Protein_CDKN2A_PDB_1a5e.png 2x" data-file-width="692" data-file-height="379" /></a></span></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd">Available structures</th></tr><tr><th rowspan="2" style="background-color:#c3fdb8;width:43px"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td colspan="2" style="background-color:#eee">Ortholog search: <span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/searchResults.html?display=both&term=P51480%20or%20Q64364%20or%20P42771%20or%20Q8N726">PDBe</a> <a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_container_identifiers.reference_sequence_identifiers.database_name:UniProt%20AND%20rcsb_polymer_entity_container_identifiers.reference_sequence_identifiers.database_accession:P51480,Q64364,P42771,Q8N726">RCSB</a> </span></td></tr><tr><td><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr style="background-color:#ddd;text-align:center"><th colspan="2">List of PDB id codes</th></tr><tr><td colspan="2" style="background-color:#eee"><p><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/2A5E">2A5E</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1A5E">1A5E</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1BI7">1BI7</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1DC2">1DC2</a>,%%s<a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1A5E">1A5E</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1BI7">1BI7</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1DC2">1DC2</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/2A5E">2A5E</a></span></p></td></tr></tbody></table></td></tr></tbody></table></td></tr><tr><th colspan="4" style="text-align:center;background-color:#ddd">Identifiers</th></tr><tr><th scope="row" style="background-color:#c3fdb8"><span class="plainlinks"><a href="/wiki/Gene_nomenclature" title="Gene nomenclature">Aliases</a></span></th><td colspan="3" style="background:#eee"><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.genenames.org/data/gene-symbol-report/#!/hgnc_id/1787">CDKN2A</a></span>, CDK4I, CDKN2, CMM2, INK4, INK4A, MLM, MTS-1, MTS1, P14, P14ARF, P16, P16-INK4A, P16INK4, P16INK4A, P19, P19ARF, TP16, cyclin-dependent kinase inhibitor 2A, cyclin dependent kinase inhibitor 2A, Genes, p16, ARF.</td></tr><tr><th scope="row" style="background-color:#c3fdb8">External IDs</th><td colspan="3" style="background-color:#eee"><span class="plainlinks"><a href="/wiki/Mendelian_Inheritance_in_Man" class="mw-redirect" title="Mendelian Inheritance in Man">OMIM</a>: <a rel="nofollow" class="external text" href="https://omim.org/entry/600160">600160</a>; <a href="/wiki/Mouse_Genome_Informatics" title="Mouse Genome Informatics">MGI</a>: <a rel="nofollow" class="external text" href="http://www.informatics.jax.org/marker/MGI:104738">104738</a>; <a href="/wiki/HomoloGene" title="HomoloGene">HomoloGene</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=homologene&dopt=HomoloGene&list_uids=55430">55430</a>; <a href="/wiki/GeneCards" title="GeneCards">GeneCards</a>: <a rel="nofollow" class="external text" href="https://www.genecards.org/cgi-bin/carddisp.pl?gene=CDKN2A">CDKN2A</a>; <a href="/wiki/Orthologous_MAtrix" title="Orthologous MAtrix">OMA</a>:<a rel="nofollow" class="external text" href="https://omabrowser.org/oma/vps/ENSG00000147889">CDKN2A - orthologs</a></span></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd">Gene location (<a href="/wiki/Human_genome" title="Human genome">Human</a>)</th></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><span typeof="mw:File"><a href="/wiki/File:Ideogram_human_chromosome_9.svg" class="mw-file-description" title="Chromosome 9 (human)"><img alt="Chromosome 9 (human)" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Ideogram_human_chromosome_9.svg/300px-Ideogram_human_chromosome_9.svg.png" decoding="async" width="300" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Ideogram_human_chromosome_9.svg/450px-Ideogram_human_chromosome_9.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/de/Ideogram_human_chromosome_9.svg/600px-Ideogram_human_chromosome_9.svg.png 2x" data-file-width="474" data-file-height="189" /></a></span></td></tr><tr><th scope="row" width="15%" style="background-color:#c3fdb8"><a href="/wiki/Chromosome" title="Chromosome">Chr.</a></th><td colspan="3" width="85%" style="background-color:#eee"><span class="plainlinks"><a href="/wiki/Chromosome_9_(human)" class="mw-redirect" title="Chromosome 9 (human)">Chromosome 9 (human)</a><sup id="cite_ref-refGRCh38Ensembl_1-0" class="reference"><a href="#cite_note-refGRCh38Ensembl-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></span></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><div align="center"><div style="position: relative; width: 300px;"><span typeof="mw:File"><a href="/wiki/File:Human_chromosome_9_ideogram.svg" class="mw-file-description" title="Chromosome 9 (human)"><img alt="Chromosome 9 (human)" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Human_chromosome_9_ideogram.svg/300px-Human_chromosome_9_ideogram.svg.png" decoding="async" width="300" height="58" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Human_chromosome_9_ideogram.svg/450px-Human_chromosome_9_ideogram.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Human_chromosome_9_ideogram.svg/600px-Human_chromosome_9_ideogram.svg.png 2x" data-file-width="1125" data-file-height="216" /></a></span><div style="position: absolute; left: 41.31780014789px; top: 2px; padding: 0;"><span typeof="mw:File"><a href="/wiki/File:HSR_1996_II_3.5e.svg" class="mw-file-description" title="Genomic location for CDKN2A"><img alt="Genomic location for CDKN2A" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/14px-HSR_1996_II_3.5e.svg.png" decoding="async" width="14" height="14" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/21px-HSR_1996_II_3.5e.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/28px-HSR_1996_II_3.5e.svg.png 2x" data-file-width="142" data-file-height="142" /></a></span></div><div style="position: absolute; left: 47.3px; top: 19px; padding: 0;"><span typeof="mw:File"><a href="/wiki/File:Red_rectangle_2x18.png" class="mw-file-description" title="Genomic location for CDKN2A"><img alt="Genomic location for CDKN2A" src="//upload.wikimedia.org/wikipedia/commons/6/6a/Red_rectangle_2x18.png" decoding="async" width="2" height="18" class="mw-file-element" data-file-width="2" data-file-height="18" /></a></span></div></div></div></td></tr><tr><th scope="row" rowspan="2" width="15%" style="background-color:#c3fdb8"><a href="/wiki/Locus_(genetics)" title="Locus (genetics)">Band</a></th><td rowspan="2" width="35%" style="background-color:#eee"><span class="plainlinks">9p21.3</span></td><th scope="row" style="background-color:#c3fdb8">Start</th><td style="background-color:#eee"><span class="plainlinks">21,967,752 <a href="/wiki/Base_pair" title="Base pair">bp</a><sup id="cite_ref-refGRCh38Ensembl_1-1" class="reference"><a href="#cite_note-refGRCh38Ensembl-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></span></td></tr><tr><th scope="row" style="background-color:#c3fdb8">End</th><td style="background-color:#eee"><span class="plainlinks">21,995,301 <a href="/wiki/Base_pair" title="Base pair">bp</a><sup id="cite_ref-refGRCh38Ensembl_1-2" class="reference"><a href="#cite_note-refGRCh38Ensembl-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></span></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd">Gene location (<a href="/wiki/Laboratory_mouse" title="Laboratory mouse">Mouse</a>)</th></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><span typeof="mw:File"><a href="/wiki/File:Ideogram_house_mouse_chromosome_4.svg" class="mw-file-description" title="Chromosome 4 (mouse)"><img alt="Chromosome 4 (mouse)" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Ideogram_house_mouse_chromosome_4.svg/260px-Ideogram_house_mouse_chromosome_4.svg.png" decoding="async" width="260" height="111" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Ideogram_house_mouse_chromosome_4.svg/390px-Ideogram_house_mouse_chromosome_4.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Ideogram_house_mouse_chromosome_4.svg/520px-Ideogram_house_mouse_chromosome_4.svg.png 2x" data-file-width="470" data-file-height="200" /></a></span></td></tr><tr><th scope="row" width="15%" style="background-color:#c3fdb8"><a href="/wiki/Chromosome" title="Chromosome">Chr.</a></th><td colspan="3" width="85%" style="background-color:#eee"><span class="plainlinks">Chromosome 4 (mouse)<sup id="cite_ref-refGRCm38Ensembl_2-0" class="reference"><a href="#cite_note-refGRCm38Ensembl-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></span></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><div align="center"><div style="position: relative; width: 300px;"><span typeof="mw:File"><a href="/wiki/File:Ideogram_of_house_mouse_chromosome_4.svg" class="mw-file-description" title="Chromosome 4 (mouse)"><img alt="Chromosome 4 (mouse)" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Ideogram_of_house_mouse_chromosome_4.svg/300px-Ideogram_of_house_mouse_chromosome_4.svg.png" decoding="async" width="300" height="58" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Ideogram_of_house_mouse_chromosome_4.svg/450px-Ideogram_of_house_mouse_chromosome_4.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Ideogram_of_house_mouse_chromosome_4.svg/600px-Ideogram_of_house_mouse_chromosome_4.svg.png 2x" data-file-width="1200" data-file-height="231" /></a></span><div style="position: absolute; left: 162.24309346633px; top: 2px; padding: 0;"><span typeof="mw:File"><a href="/wiki/File:HSR_1996_II_3.5e.svg" class="mw-file-description" title="Genomic location for CDKN2A"><img alt="Genomic location for CDKN2A" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/14px-HSR_1996_II_3.5e.svg.png" decoding="async" width="14" height="14" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/21px-HSR_1996_II_3.5e.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/HSR_1996_II_3.5e.svg/28px-HSR_1996_II_3.5e.svg.png 2x" data-file-width="142" data-file-height="142" /></a></span></div><div style="position: absolute; left: 168.2px; top: 19px; padding: 0;"><span typeof="mw:File"><a href="/wiki/File:Red_rectangle_2x18.png" class="mw-file-description" title="Genomic location for CDKN2A"><img alt="Genomic location for CDKN2A" src="//upload.wikimedia.org/wikipedia/commons/6/6a/Red_rectangle_2x18.png" decoding="async" width="2" height="18" class="mw-file-element" data-file-width="2" data-file-height="18" /></a></span></div></div></div></td></tr><tr><th scope="row" rowspan="2" width="15%" style="background-color:#c3fdb8"><a href="/wiki/Locus_(genetics)" title="Locus (genetics)">Band</a></th><td rowspan="2" width="35%" style="background-color:#eee"><span class="plainlinks">4 C4|4 42.15 cM</span></td><th scope="row" style="background-color:#c3fdb8">Start</th><td style="background-color:#eee"><span class="plainlinks">89,192,708 <a href="/wiki/Base_pair" title="Base pair">bp</a><sup id="cite_ref-refGRCm38Ensembl_2-1" class="reference"><a href="#cite_note-refGRCm38Ensembl-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></span></td></tr><tr><th scope="row" style="background-color:#c3fdb8">End</th><td style="background-color:#eee"><span class="plainlinks">89,212,890 <a href="/wiki/Base_pair" title="Base pair">bp</a><sup id="cite_ref-refGRCm38Ensembl_2-2" class="reference"><a href="#cite_note-refGRCm38Ensembl-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></span></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd"><a href="/wiki/Gene_expression" title="Gene expression">RNA expression</a> pattern</th></tr><tr><th scope="row" style="background-color:#c3fdb8"><a rel="nofollow" class="external text" href="https://www.bgee.org/">Bgee</a></th><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th><b><a href="/wiki/Human_genome" title="Human genome">Human</a></b></th><th><b><a href="/wiki/Laboratory_mouse" title="Laboratory mouse">Mouse</a> (ortholog)</b></th></tr><tr><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:center"><tbody><tr><td colspan="1"><span class="plainlinks" style="margin:-3px"><a rel="nofollow" class="external text" href="https://www.bgee.org/gene/ENSG00000147889">Top expressed in</a></span></td></tr><tr><td colspan="1"><div class="plainlinks" style="margin:-12px 0px -10px 0px"><ul style="line-height:15%;margin:9px"><li style="line-height: 137%;">parotid gland</li><br /><li style="line-height: 137%;">pituitary gland</li><br /><li style="line-height: 137%;">testicle</li><br /><li style="line-height: 137%;">anterior pituitary</li><br /><li style="line-height: 137%;">subthalamic nucleus</li><br /><li style="line-height: 137%;">tongue</li><br /><li style="line-height: 137%;">ventral tegmental area</li><br /><li style="line-height: 137%;">mucosa of pharynx</li><br /><li style="line-height: 137%;">superior surface of tongue</li><br /><li style="line-height: 137%;">external globus pallidus</li></ul></div></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:center"><tbody><tr><td colspan="1"><span class="plainlinks" style="margin:-3px"><a rel="nofollow" class="external text" href="https://www.bgee.org/gene/ENSMUSG00000044303">Top expressed in</a></span></td></tr><tr><td colspan="1"><div class="plainlinks" style="margin:-12px 0px -10px 0px"><ul style="line-height:15%;margin:9px"><li style="line-height: 137%;">yolk sac</li><br /><li style="line-height: 137%;">stroma of bone marrow</li><br /><li style="line-height: 137%;">morula</li><br /><li style="line-height: 137%;">embryo</li><br /><li style="line-height: 137%;">glossopharyngeal ganglion</li><br /><li style="line-height: 137%;">endothelial cell of lymphatic vessel</li><br /><li style="line-height: 137%;">lumbar subsegment of spinal cord</li><br /><li style="line-height: 137%;">calvaria</li><br /><li style="line-height: 137%;">greater petrosal nerve</li><br /><li style="line-height: 137%;">muscle of thigh</li></ul></div></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.bgee.org/gene/ENSG00000147889">More reference expression data</a></span></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8"><a rel="nofollow" class="external text" href="http://biogps.org/">BioGPS</a></th><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><td colspan="4" style="text-align:center;background-color:#eee"><span typeof="mw:File"><a href="/wiki/File:PBB_GE_CDKN2A_207039_at.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/PBB_GE_CDKN2A_207039_at.png/250px-PBB_GE_CDKN2A_207039_at.png" decoding="async" width="250" height="132" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/e6/PBB_GE_CDKN2A_207039_at.png 1.5x" data-file-width="255" data-file-height="135" /></a></span><br /><br /><span typeof="mw:File"><a href="/wiki/File:PBB_GE_CDKN2A_209644_x_at.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a1/PBB_GE_CDKN2A_209644_x_at.png/250px-PBB_GE_CDKN2A_209644_x_at.png" decoding="async" width="250" height="132" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/a/a1/PBB_GE_CDKN2A_209644_x_at.png 1.5x" data-file-width="255" data-file-height="135" /></a></span><br /><br /><span typeof="mw:File"><a href="/wiki/File:PBB_GE_CDKN2A_211156_at.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/PBB_GE_CDKN2A_211156_at.png/250px-PBB_GE_CDKN2A_211156_at.png" decoding="async" width="250" height="132" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/1/10/PBB_GE_CDKN2A_211156_at.png 1.5x" data-file-width="255" data-file-height="135" /></a></span></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><span class="plainlinks"><a rel="nofollow" class="external text" href="http://biogps.org/gene/1029/">More reference expression data</a></span></td></tr></tbody></table></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd"><a href="/wiki/Gene_ontology" class="mw-redirect" title="Gene ontology">Gene ontology</a></th></tr><tr><td style="background-color:#c3fdb8;font-weight:bold">Molecular function</td><td style="background-color:#eee"><div class="plainlinks"> <ul><li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0051059">NF-kappaB binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005515">protein binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0004861">cyclin-dependent protein serine/threonine kinase inhibitor activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0019901">protein kinase binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0003723">RNA binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0003677">DNA binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0002039">p53 binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0008134">transcription factor binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0097371">MDM2/MDM4 family protein binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0055105">ubiquitin-protein transferase inhibitor activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0019789">SUMO transferase activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0097718">disordered domain specific binding</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1990948">ubiquitin ligase inhibitor activity</a></li></ul> </div></td></tr><tr><td style="background-color:#c3fdb8;font-weight:bold">Cellular component</td><td style="background-color:#eee"><div class="plainlinks"> <ul><li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005737">cytoplasm</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005829">cytosol</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0035985">senescence-associated heterochromatin focus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005634">nucleus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0016604">nuclear body</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005654">nucleoplasm</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005730">nucleolus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0005739">mitochondrion</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0032991">protein-containing complex</a></li></ul> </div></td></tr><tr><td style="background-color:#c3fdb8;font-weight:bold">Biological process</td><td style="background-color:#eee"><div class="plainlinks"> <ul><li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0034393">positive regulation of smooth muscle cell apoptotic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0045736">negative regulation of cyclin-dependent protein serine/threonine kinase activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0090399">replicative senescence</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0051318">G1 phase</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0001953">negative regulation of cell-matrix adhesion</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0090398">cellular senescence</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0030308">negative regulation of cell growth</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:2000774">positive regulation of cellular senescence</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:2000111">positive regulation of macrophage apoptotic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0007049">cell cycle</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0007265">Ras protein signal transduction</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0042326">negative regulation of phosphorylation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0045892">negative regulation of transcription, DNA-templated</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0032088">negative regulation of NF-kappaB transcription factor activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0008285">negative regulation of cell population proliferation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0000082">G1/S transition of mitotic cell cycle</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:2000134">negative regulation of G1/S transition of mitotic cell cycle</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006915">apoptotic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0010389">regulation of G2/M transition of mitotic cell cycle</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006355">regulation of transcription, DNA-templated</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0031647">regulation of protein stability</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0033088">negative regulation of immature T cell proliferation in thymus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0046825">regulation of protein export from nucleus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006469">negative regulation of protein kinase activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1903051">negative regulation of proteolysis involved in cellular protein catabolic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0070534">protein K63-linked ubiquitination</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1901798">positive regulation of signal transduction by p53 class mediator</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0048103">somatic stem cell division</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0050821">protein stabilization</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0000209">protein polyubiquitination</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0033235">positive regulation of protein sumoylation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006351">transcription, DNA-templated</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0045893">positive regulation of transcription, DNA-templated</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0000422">autophagy of mitochondrion</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0008637">apoptotic mitochondrial changes</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1903214">regulation of protein targeting to mitochondrion</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0031648">protein destabilization</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0051444">negative regulation of ubiquitin-protein transferase activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0043065">positive regulation of apoptotic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006364">rRNA processing</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:2000059">negative regulation of ubiquitin-dependent protein catabolic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0051882">mitochondrial depolarization</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0043517">positive regulation of DNA damage response, signal transduction by p53 class mediator</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0030889">negative regulation of B cell proliferation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0006919">activation of cysteine-type endopeptidase activity involved in apoptotic process</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1902510">regulation of apoptotic DNA fragmentation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0045944">positive regulation of transcription by RNA polymerase II</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1900182">positive regulation of protein localization to nucleus</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0016925">protein sumoylation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0010628">positive regulation of gene expression</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1904667">negative regulation of ubiquitin protein ligase activity</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:1990000">amyloid fibril formation</a></li> <li><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:2000435">negative regulation of protein neddylation</a></li></ul> </div></td></tr><tr><td colspan="4" style="background-color:#eee;text-align:center">Sources:<a rel="nofollow" class="external text" href="http://amigo.geneontology.org/">Amigo</a> / <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/QuickGO/">QuickGO</a></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table class="collapsible" style="padding:0;border:none;margin:0;width:100%;text-align:left"><tbody><tr><th colspan="4" style="text-align:center;background-color:#ddd"><a href="/wiki/Orthologs" class="mw-redirect" title="Orthologs">Orthologs</a></th></tr><tr><th scope="row" style="background-color:#c3fdb8">Species</th><td><b>Human</b></td><td><b>Mouse</b></td></tr><tr><th scope="row" style="background-color:#c3fdb8"><a href="/wiki/Entrez" title="Entrez">Entrez</a></th><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene&cmd=retrieve&dopt=default&list_uids=1029&rn=1">1029</a></p></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene&cmd=retrieve&dopt=default&list_uids=12578&rn=1">12578</a></p></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8"><a href="/wiki/Ensembl" class="mw-redirect" title="Ensembl">Ensembl</a></th><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="http://www.ensembl.org/Homo_sapiens/geneview?gene=ENSG00000147889;db=core">ENSG00000147889</a></p></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="http://www.ensembl.org/Mus_musculus/geneview?gene=ENSMUSG00000044303;db=core">ENSMUSG00000044303</a></p></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8"><a href="/wiki/UniProt" title="UniProt">UniProt</a></th><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P42771">P42771</a><br /><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/Q8N726">Q8N726</a></p></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1"><span class="plainlinks"></span></th></tr><tr><td colspan="1"><p class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P51480">P51480</a><br /><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/Q64364">Q64364</a></p></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8">RefSeq (mRNA)</th><td><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1" class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_000077">NM_000077</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_001195132">NM_001195132</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_058195">NM_058195</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_058196">NM_058196</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_058197">NM_058197</a><br /></th></tr><tr><td colspan="1"><p><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_001363763">NM_001363763</a></span></p></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1" class="plainlinks"></th></tr><tr><td colspan="1"><p><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_001040654">NM_001040654</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_009877">NM_009877</a></span></p></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8">RefSeq (protein)</th><td><table class="collapsible collapsed" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1" class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_000068">NP_000068</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_001182061">NP_001182061</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_478102">NP_478102</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_478104">NP_478104</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_001350692">NP_001350692</a><br /></th></tr><tr><td colspan="1"><p><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_478102.2">NP_478102.2</a></span></p></td></tr></tbody></table></td><td><table class="none" style="padding:0;border:none;margin:0;width:100%;text-align:right"><tbody><tr><th colspan="1" class="plainlinks"></th></tr><tr><td colspan="1"><p><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_001035744">NP_001035744</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_034007">NP_034007</a><br /><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_034007.1">NP_034007.1</a></span></p></td></tr></tbody></table></td></tr><tr><th scope="row" style="background-color:#c3fdb8">Location (UCSC)</th><td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgTracks?org=Human&db=hg38&position=chr9:21967752-21995301">Chr 9: 21.97 – 22 Mb</a></span></td><td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgTracks?org=Mouse&db=mm0&position=chr4:89192708-89212890">Chr 4: 89.19 – 89.21 Mb</a></span></td></tr><tr><th scope="row" style="background-color:#c3fdb8"><a href="/wiki/PubMed" title="PubMed">PubMed</a> search</th><td><span class="plainlinks"><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></span></td><td><span class="plainlinks"><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup></span></td></tr></tbody></table></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a></td></tr><tr><td colspan="4" style="text-align:center;background-color:#eee"><table style="padding:0;border:none;margin:0;width:100%;text-align:center"><tbody><tr><td colspan="2" style="background-color:#eee;text-align:center"><a href="https://www.wikidata.org/wiki/Q5009957" class="extiw" title="d:Q5009957">View/Edit Human</a></td><td colspan="2" style="background-color:#eee;text-align:center"><a href="https://www.wikidata.org/wiki/Q14907011" class="extiw" title="d:Q14907011">View/Edit Mouse</a></td></tr></tbody></table></td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">Cyclin-dependent kinase inhibitor 2a p19Arf N-terminus</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:PDB_1hn3_EBI.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/PDB_1hn3_EBI.jpg/220px-PDB_1hn3_EBI.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/PDB_1hn3_EBI.jpg/330px-PDB_1hn3_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0a/PDB_1hn3_EBI.jpg/440px-PDB_1hn3_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a></span><div class="infobox-caption">Solution structure of the n-terminal 37 amino acids of the mouse arf tumor suppressor protein</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3">Symbol</th><td class="infobox-data" style="background-color: #eee">P19Arf_N</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF07392">PF07392</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR010868">IPR010868</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Structural_Classification_of_Proteins" class="mw-redirect" title="Structural Classification of Proteins">SCOP2</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=1hn3">1hn3</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=1hn3">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=1hn3">SUPFAM</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Pfam" title="Pfam">Pfam</a>  </th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF07392?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF07392">ECOD</a>  </td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF07392%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF07392">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF07392">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PDBsum" title="PDBsum">PDBsum</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF07392">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p><b>p16</b> (also known as <b>p16<sup>INK4a</sup></b>, <b>cyclin-dependent kinase inhibitor 2A</b>, <b>CDKN2A</b>, <b>multiple tumor suppressor 1</b> and numerous other synonyms), is a <a href="/wiki/Protein" title="Protein">protein</a> that slows <a href="/wiki/Cell_division" title="Cell division">cell division</a> by slowing the progression of the cell cycle from the <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a> to the <a href="/wiki/S_phase" title="S phase">S phase</a>, thereby acting as a <a href="/wiki/Tumor_suppressor" class="mw-redirect" title="Tumor suppressor">tumor suppressor</a>. It is encoded by the <i><a href="/wiki/CDKN2A" title="CDKN2A">CDKN2A</a></i> <a href="/wiki/Gene" title="Gene">gene</a>. A <a href="/wiki/Genetics" title="Genetics">deletion</a> (the omission of a part of the DNA sequence during replication) in this gene can result in insufficient or non-functional p16, accelerating the cell cycle and resulting in many types of cancer.<sup id="cite_ref-entrez_5-0" class="reference"><a href="#cite_note-entrez-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid8152487_6-0" class="reference"><a href="#cite_note-pmid8152487-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7606716_7-0" class="reference"><a href="#cite_note-pmid7606716-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>p16 can be used as a <a href="/wiki/Biomarker" title="Biomarker">biomarker</a> to improve the histological diagnostic accuracy of grade 3 <a href="/wiki/Cervical_intraepithelial_neoplasia" title="Cervical intraepithelial neoplasia">cervical intraepithelial neoplasia</a> (CIN). p16 is also implicated in the prevention of <a href="/wiki/Melanoma" title="Melanoma">melanoma</a>, <a href="/wiki/Oropharyngeal_cancer" title="Oropharyngeal cancer">oropharyngeal squamous cell carcinoma</a>, <a href="/wiki/Cervical_cancer" title="Cervical cancer">cervical cancer</a>, <a href="/wiki/Vulvar_cancer" title="Vulvar cancer">vulvar cancer</a> and <a href="/wiki/Esophageal_cancer" title="Esophageal cancer">esophageal cancer</a>. </p><p>p16 was discovered in 1993. It is a protein with 148 amino acids and a <a href="/wiki/Molecular_weight" class="mw-redirect" title="Molecular weight">molecular weight</a> of 16 <a href="/wiki/Kilodalton" class="mw-redirect" title="Kilodalton">kDa</a> that comprises four <a href="/wiki/Ankyrin_repeat" title="Ankyrin repeat">ankyrin repeats</a>.<sup id="cite_ref-pmid8259215_8-0" class="reference"><a href="#cite_note-pmid8259215-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The name of p16 is derived from its <a href="/wiki/Molecular_weight" class="mw-redirect" title="Molecular weight">molecular weight</a>, and the alternative name p16<sup>INK4a</sup> refers to its role in inhibiting <a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">cyclin-dependent kinase</a> CDK4.<sup id="cite_ref-pmid8259215_8-1" class="reference"><a href="#cite_note-pmid8259215-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Nomenclature">Nomenclature</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=1" title="Edit section: Nomenclature"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>p16 is also known as: </p> <ul><li>p16<sup>INK4A</sup></li> <li>p16<sup>Ink4</sup></li> <li>Cyclin-dependent kinase inhibitor 2A (CDKN2A)</li> <li>CDKN2</li> <li>CDK 4 Inhibitor</li> <li>Multiple Tumor Suppressor 1 (MTS1)</li> <li>TP16</li> <li>ARF</li> <li>MLM</li> <li>P14</li></ul> <div class="mw-heading mw-heading2"><h2 id="Gene">Gene</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=2" title="Edit section: Gene"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In humans, p16 is encoded by the <a href="/wiki/CDKN2A" title="CDKN2A">CDKN2A</a> gene, located on <a href="/wiki/Chromosome_9_(human)" class="mw-redirect" title="Chromosome 9 (human)">chromosome 9</a> (9p21.3). This <a href="/wiki/Gene" title="Gene">gene</a> generates several <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcript</a> variants that differ in their first <a href="/wiki/Exon" title="Exon">exons</a>. At least three alternatively <a href="/wiki/Splicing_(genetics)" class="mw-redirect" title="Splicing (genetics)">spliced</a> variants encoding distinct proteins have been reported, two of which encode structurally related <a href="/wiki/Protein_isoform" title="Protein isoform">isoforms</a> known to function as <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">inhibitors</a> of <a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4</a>. The remaining transcript includes an alternate exon 1 located 20 <a href="/wiki/Kilobase" class="mw-redirect" title="Kilobase">kb</a> upstream of the remainder of the gene; this transcript contains an <a href="/wiki/P14arf" title="P14arf">alternate open reading frame (ARF)</a> that specifies a protein that is structurally unrelated to the products of the other variants.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> The ARF product functions as a stabilizer of the tumor suppressor protein <a href="/wiki/P53_(protein)" class="mw-redirect" title="P53 (protein)">p53</a>, as it can interact with and sequester <a href="/wiki/MDM2" class="mw-redirect" title="MDM2">MDM2</a>, a protein responsible for the degradation of p53.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10498883_11-0" class="reference"><a href="#cite_note-pmid10498883-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> In spite of their structural and functional differences, the CDK inhibitor isoforms and the ARF product encoded by this gene, through the regulatory roles of CDK4 and p53 in <a href="/wiki/Cell_cycle#G1_phase" title="Cell cycle">cell cycle G1 progression</a>, share a common functionality in controlling the G1 phase of the cell cycle. This gene is frequently mutated or deleted in a wide variety of tumors and is known to be an important tumor suppressor gene.<sup id="cite_ref-entrez_5-1" class="reference"><a href="#cite_note-entrez-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>When organisms age, the expression of p16 increases to reduce the proliferation of <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a>.<sup id="cite_ref-pmid16957737_12-0" class="reference"><a href="#cite_note-pmid16957737-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> This reduction in the division and production of stem cells protects against <a href="/wiki/Cancer" title="Cancer">cancer</a> while increasing the risks associated with <a href="/wiki/Senescence" title="Senescence">cellular senescence</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=3" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>p16 is an inhibitor of cyclin-dependent kinases (CDK). It slows down the cell cycle by prohibiting progression from G1 phase to S phase. Otherwise, CDK4/6 binds cyclin D and forms an active protein complex that phosphorylates <a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">retinoblastoma protein</a> (pRB). Once phosphorylated, pRB dissociates from the transcription factor <a href="/wiki/E2F1" title="E2F1">E2F1</a>. This liberates E2F1 from its bound state in the cytoplasm and allows it to enter the nucleus. Once in the nucleus, E2F1 promotes the transcription of target genes that are essential for transition from G1 to S phase.<sup id="cite_ref-Rayess2012_13-0" class="reference"><a href="#cite_note-Rayess2012-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid8622687_14-0" class="reference"><a href="#cite_note-pmid8622687-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>This pathway connects the processes of tumor oncogenesis and senescence, fixing them on opposite ends of a spectrum. On one end, p16 hypermethylation, mutation, or deletion leads to downregulation of the gene and can lead to cancer through the dysregulation of cell cycle progression. Conversely, activation of p16 through <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a>, DNA damage, or senescence leads to the buildup of p16 in tissues and is implicated in the aging of cells.<sup id="cite_ref-Rayess2012_13-1" class="reference"><a href="#cite_note-Rayess2012-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Regulation">Regulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=4" title="Edit section: Regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Regulation of p16 is complex and involves the interaction of several transcription factors, as well as several proteins involved in epigenetic modification through methylation and repression of the promoter region.<sup id="cite_ref-Rayess2012_13-2" class="reference"><a href="#cite_note-Rayess2012-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>PRC1 and PRC2 are two protein complexes that modify the expression of p16 through the interaction of various transcription factors that execute methylation patterns that can repress transcription of p16. These pathways are activated in the cellular response to reduce senescence.<sup id="cite_ref-pmid12351676_15-0" class="reference"><a href="#cite_note-pmid12351676-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid17344414_16-0" class="reference"><a href="#cite_note-pmid17344414-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_significance">Clinical significance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=5" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Role_in_carcinogenesis">Role in carcinogenesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=6" title="Edit section: Role in carcinogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mutations resulting in deletion or reduction of function of the CDKN2A gene are associated with increased risk of a wide range of cancers, and alterations of the gene are frequently seen in cancer <a href="/wiki/Cell_line" class="mw-redirect" title="Cell line">cell lines</a>.<sup id="cite_ref-pmid9508208_17-0" class="reference"><a href="#cite_note-pmid9508208-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11237522_18-0" class="reference"><a href="#cite_note-pmid11237522-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Examples include: </p><p><a href="/wiki/Pancreatic_cancer" title="Pancreatic cancer">Pancreatic adenocarcinoma</a> is often associated with mutations in the CDKN2A gene.<sup id="cite_ref-pmid7726912_19-0" class="reference"><a href="#cite_note-pmid7726912-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid8589035_20-0" class="reference"><a href="#cite_note-pmid8589035-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7624155_21-0" class="reference"><a href="#cite_note-pmid7624155-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Carriers of <a href="/wiki/Germline_mutation" title="Germline mutation">germline mutations</a> in CDKN2A have, besides their high risks of melanoma, also increased risks of pancreatic, lung, laryngeal and oropharyngeal cancers. Tobacco smoking increases the carriers’ susceptibility for such non-melanoma cancers.<sup id="cite_ref-PMID24935963_22-0" class="reference"><a href="#cite_note-PMID24935963-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>Homozygous deletions of p16 are frequently found in <a href="/wiki/Esophageal_cancer" title="Esophageal cancer">esophageal cancer</a> and <a href="/wiki/Gastric_cancer" class="mw-redirect" title="Gastric cancer">gastric cancer</a> cell lines.<sup id="cite_ref-pmid8093026_23-0" class="reference"><a href="#cite_note-pmid8093026-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>Germline mutations in CDKN2A are associated with an increased susceptibility to develop <a href="/wiki/Skin_cancer" title="Skin cancer">skin cancer</a>.<sup id="cite_ref-pmid24742402_24-0" class="reference"><a href="#cite_note-pmid24742402-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Hypermethylation of tumor suppressor genes has been implicated in various cancers. In 2013, a meta-analysis revealed an increased frequency of DNA methylation of the p16 gene in esophageal cancer. As the degree of tumor differentiation increased, so did the frequency of p16 DNA methylation. </p><p>Tissue samples of primary oral squamous cell carcinoma (OSCC) often display hypermethylation in the promoter regions of p16. Cancer cells show a significant increase in the accumulation of methylation in CpG islands in the promoter region of p16. This <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> change leads to loss of the tumor suppressor gene function through two possible mechanisms: first, methylation can physically inhibit the transcription of the gene, and second, methylation can lead to the recruitment of transcription factors that repress transcription. Both mechanisms cause the same end result: downregulation of gene expression that leads to decreased levels of the p16 protein. It has been suggested that this process is responsible for the development of various forms of cancer serving as an alternative process to gene deletion or mutation.<sup id="cite_ref-pmid24155659_25-0" class="reference"><a href="#cite_note-pmid24155659-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid22106032_26-0" class="reference"><a href="#cite_note-pmid22106032-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid16449996_27-0" class="reference"><a href="#cite_note-pmid16449996-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid17383681_28-0" class="reference"><a href="#cite_note-pmid17383681-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid22295037_29-0" class="reference"><a href="#cite_note-pmid22295037-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24240582_30-0" class="reference"><a href="#cite_note-pmid24240582-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>p16 positivity has been shown to be favorably prognostic in oropharyngeal squamous cell carcinoma.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> In a retrospective trial analysis of patients with Stage III and IV oropharyngeal cancer, HPV status was assessed and it was found that the 3-year rates of overall survival were 82.4% (95% CI, 77.2 to 87.6) in the HPV-positive subgroup and 57.1% (95% CI, 48.1 to 66.1) in the HPV-negative subgroup, and the 3-year rates of progression-free survival were 73.7% (95% CI, 67.7 to 79.8) and 43.4% (95% CI, 34.4 to 52.4), respectively. p16 status is so prognostic that the AJCC staging system has been revised to include p16 status in oropharyngeal squamous cell cancer group staging.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> However, some people can have elevated levels of p16 but test negative for HPV and vice versa. This is known as discordant cancer. The 5-year survival for people who test positive for HPV and p16 is 81%, for discordant cancer it is 53–55%, and 40% for those who test negative for p16 and HPV.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_use">Clinical use</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=7" title="Edit section: Clinical use"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Biomarker_for_cancer_types">Biomarker for cancer types</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=8" title="Edit section: Biomarker for cancer types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Expression of p16 is used as a prognostic biomarker for certain types of cancer. The reason for this is different types of cancer can have different effects on p16 expression: cancers that overexpress p16 are usually caused by the <a href="/wiki/Human_papillomavirus" class="mw-redirect" title="Human papillomavirus">human papillomavirus</a> (HPV), whereas cancers in which p16 is downregulated will usually have other causes. For patients with oropharyngeal squamous cell carcinoma, using immunohistochemistry to detect the presence of the p16 biomarker has been shown to be the strongest indicator of disease course. Presence of the biomarker is associated with a more favorable prognosis as measured by cancer-specific survival (CSS), recurrence-free survival (RFS), locoregional control (LRC), as well as other measurements. The appearance of hypermethylation of p16 is also being evaluated as a potential prognostic biomarker for prostate cancer.<sup id="cite_ref-pmid23916365_35-0" class="reference"><a href="#cite_note-pmid23916365-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid23387484_36-0" class="reference"><a href="#cite_note-pmid23387484-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24211084_37-0" class="reference"><a href="#cite_note-pmid24211084-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="p16_FISH">p16 FISH</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=9" title="Edit section: p16 FISH"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>p16 <a href="/wiki/Deletion_(genetics)" title="Deletion (genetics)">deletion</a> detected by <a href="/wiki/FISH_(genetic)" class="mw-redirect" title="FISH (genetic)">FISH</a> in surface epithelial mesothelial proliferations is predictive of underlying invasive <a href="/wiki/Mesothelioma" title="Mesothelioma">mesothelioma</a>.<sup id="cite_ref-pmid24503757_38-0" class="reference"><a href="#cite_note-pmid24503757-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="p16_immunochemistry">p16 immunochemistry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=10" title="Edit section: p16 immunochemistry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg/220px-High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg/330px-High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/92/High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg/440px-High_grade_squamous_intraepithelial_lesion_-_2_-_p16_--_high_mag.jpg 2x" data-file-width="4272" data-file-height="2848" /></a><figcaption>High grade squamous intraepithelial lesion showing strong p16 staining</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg/220px-Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg/330px-Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/66/Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg/440px-Immunohistochemistry_for_p16_in_uterine_papillary_serous_adenocarcinoma_showing_both_nuclear_and_cytoplasmic_staining.jpg 2x" data-file-width="2048" data-file-height="1532" /></a><figcaption>To count as positive, p16 immunohistochemistry should show "block" staining, which is strong nuclear and cytoplasmic expression in a continuous segment of cells.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>As consensus grows regarding the strength of p16 as a biomarker for detecting and determining prognoses of cancer, p16 immunohistochemistry is growing in importance.<sup id="cite_ref-Rayess2012_13-3" class="reference"><a href="#cite_note-Rayess2012-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid23916365_35-1" class="reference"><a href="#cite_note-pmid23916365-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid23503925_40-0" class="reference"><a href="#cite_note-pmid23503925-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Gynecologic_cancers">Gynecologic cancers</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=11" title="Edit section: Gynecologic cancers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>p16 is a widely used immunohistochemical marker in gynecologic pathology. Strong and diffuse cytoplasmic and nuclear expression of p16 in <a href="/wiki/Squamous_cell_carcinoma" class="mw-redirect" title="Squamous cell carcinoma">squamous cell carcinomas</a> (SCC) of the female genital tract is strongly associated with high-risk <a href="/wiki/Human_papilloma_virus" class="mw-redirect" title="Human papilloma virus">human papilloma virus</a> (HPV) infection and neoplasms of cervical origin. The majority of SCCs of uterine cervix express p16. However, p16 can be expressed in other neoplasms and in several normal human tissues.<sup id="cite_ref-pmid20571342_41-0" class="reference"><a href="#cite_note-pmid20571342-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Urinary_bladder_SCCs">Urinary bladder SCCs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=12" title="Edit section: Urinary bladder SCCs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>More than a third of urinary bladder SCCs express p16. SCCs of urinary bladder express p16 independent of gender. p16 immunohistochemical expression alone cannot be used to discriminate between SCCs arising from uterine cervix versus urinary bladder.<sup id="cite_ref-pmid20571342_41-1" class="reference"><a href="#cite_note-pmid20571342-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Role_in_cellular_senescence">Role in cellular senescence</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=13" title="Edit section: Role in cellular senescence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Concentrations of p16INK4a increase dramatically as tissue ages. p16INK4a, along with <a href="/wiki/Senescence-associated_beta-galactosidase" title="Senescence-associated beta-galactosidase">senescence-associated beta-galactosidase</a>, is regarded to be a <a href="/wiki/Biomarker" title="Biomarker">biomarker</a> of <a href="/wiki/Senescence#Cellular_senescence" title="Senescence">cellular senescence</a>.<sup id="cite_ref-pmid27391570_42-0" class="reference"><a href="#cite_note-pmid27391570-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Therefore, p16INK4a could potentially be used as a blood test that measures how fast the body's tissues are aging at a molecular level.<sup id="cite_ref-pmid19485966_43-0" class="reference"><a href="#cite_note-pmid19485966-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Notably, a recent survey of cellular senescence induced by multiple treatments to several cell lines does not identify p16 as belonging to a "core signature" of senescence markers.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>It has been used as a target to delay some aging changes in mice.<sup id="cite_ref-Baker2011_45-0" class="reference"><a href="#cite_note-Baker2011-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Role_in_neurogenesis">Role in neurogenesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=14" title="Edit section: Role in neurogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Increasing p16INK4a expression during aging is associated with reduced progenitor functions from the subventricular zone, which generates throughout life new neurons migrating to the olfactory bulb, thereby reducing olfactory neurogenesis.<sup id="cite_ref-Molofsky_et_al.,_2006_46-0" class="reference"><a href="#cite_note-Molofsky_et_al.,_2006-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Deletion of p16INK4a does not affect neurogenesis in the other adult neurogenic niche, the <a href="/wiki/Dentate_gyrus" title="Dentate gyrus">dentate gyrus</a> of the hippocampus.<sup id="cite_ref-Molofsky_et_al.,_2006_46-1" class="reference"><a href="#cite_note-Molofsky_et_al.,_2006-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> However, recently, it has been demonstrated that p16INK4a protects from depletion after a powerful proneurogenic stimulus—i.e., running— also stem and progenitor cells of the aged dentate gyrus.<sup id="cite_ref-Micheli_et_al.,_2019_47-0" class="reference"><a href="#cite_note-Micheli_et_al.,_2019-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> In fact, after deletion of p16INK4a, stem cells of the dentate gyrus are greatly activated by running, while, in wild-type p16INK4a dentate gyrus stem cells are not affected by running.<sup id="cite_ref-Micheli_et_al.,_2019_47-1" class="reference"><a href="#cite_note-Micheli_et_al.,_2019-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Therefore, p16Ink4a plays a role in the maintenance of dentate gyrus stem cells after stimulus, by keeping a reserve of their self-renewal capacity during aging. Since the dentate gyrus plays a key role in spatial and contextual memory formation, p16INK4a is implicated in the maintenance of cognitive functions during aging. </p> <div class="mw-heading mw-heading2"><h2 id="Discovery">Discovery</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=15" title="Edit section: Discovery"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Researchers Manuel Serrano, Gregory J. Hannon and David Beach discovered p16 in 1993 and correctly characterized the protein as a cyclin-dependent kinase inhibitor. </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_carcinogenesis_2">Role in carcinogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=16" title="Edit section: Role in carcinogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Since its discovery, p16 has become significant in the field of cancer research. The protein was suspected to be involved in carcinogenesis due to the observation that mutation or deletion in the gene was implicated in human cancer cell lines. The detection of p16 inactivation in familial melanoma supplied further evidence. p16 deletion, mutation, hypermethylation, or overexpression is now associated with various cancers. Whether mutations in p16 can be considered to be driver mutations requires further investigation.<sup id="cite_ref-pmid9508208_17-1" class="reference"><a href="#cite_note-pmid9508208-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Interactions">Interactions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=17" title="Edit section: Interactions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>p16 has been shown to <a href="/wiki/Protein-protein_interaction" class="mw-redirect" title="Protein-protein interaction">interact</a> with: </p> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 30em;"> <ul><li><a href="/wiki/CCNG1" title="CCNG1">CCNG1</a>,<sup id="cite_ref-pmid12556559_48-0" class="reference"><a href="#cite_note-pmid12556559-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4</a>,<sup id="cite_ref-pmid8259215_8-2" class="reference"><a href="#cite_note-pmid8259215-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid15065884_49-0" class="reference"><a href="#cite_note-pmid15065884-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10580009_50-0" class="reference"><a href="#cite_note-pmid10580009-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid17353931_51-0" class="reference"><a href="#cite_note-pmid17353931-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid8805225_52-0" class="reference"><a href="#cite_note-pmid8805225-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9228064_53-0" class="reference"><a href="#cite_note-pmid9228064-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Cyclin-dependent_kinase_6" title="Cyclin-dependent kinase 6">CDK6</a>,<sup id="cite_ref-pmid8805225_52-1" class="reference"><a href="#cite_note-pmid8805225-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9751050_54-0" class="reference"><a href="#cite_note-pmid9751050-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11739795_55-0" class="reference"><a href="#cite_note-pmid11739795-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Death_associated_protein_6" class="mw-redirect" title="Death associated protein 6">DAXX</a>,<sup id="cite_ref-pmid18583933_56-0" class="reference"><a href="#cite_note-pmid18583933-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/E4F1" title="E4F1">E4F1</a>,<sup id="cite_ref-pmid12446718_57-0" class="reference"><a href="#cite_note-pmid12446718-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Mdm2" title="Mdm2">MDM2</a>,<sup id="cite_ref-pmid18583933_56-1" class="reference"><a href="#cite_note-pmid18583933-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid14612427_58-0" class="reference"><a href="#cite_note-pmid14612427-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9529249_59-0" class="reference"><a href="#cite_note-pmid9529249-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid12085228_60-0" class="reference"><a href="#cite_note-pmid12085228-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9529248_61-0" class="reference"><a href="#cite_note-pmid9529248-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/P53" title="P53">P53</a>,<sup id="cite_ref-pmid12446718_57-1" class="reference"><a href="#cite_note-pmid12446718-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid14612427_58-1" class="reference"><a href="#cite_note-pmid14612427-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9529249_59-1" class="reference"><a href="#cite_note-pmid9529249-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/PPP1R9B" title="PPP1R9B">PPP1R9B</a>,<sup id="cite_ref-pmid11278317_62-0" class="reference"><a href="#cite_note-pmid11278317-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/RPL11" class="mw-redirect" title="RPL11">RPL11</a>,<sup id="cite_ref-pmid14612427_58-2" class="reference"><a href="#cite_note-pmid14612427-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> and</li> <li><a href="/wiki/SERTAD1" title="SERTAD1">SERTAD1</a>.<sup id="cite_ref-pmid15065884_49-1" class="reference"><a href="#cite_note-pmid15065884-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10580009_50-1" class="reference"><a href="#cite_note-pmid10580009-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/P21" title="P21">p21</a></li> <li><a href="/wiki/P53" title="P53">p53</a></li> <li><a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">Cyclin-dependent kinase</a></li> <li><a href="/wiki/Cyclin_D" title="Cyclin D">Cyclin D</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=19" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-refGRCh38Ensembl-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-refGRCh38Ensembl_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-refGRCh38Ensembl_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-refGRCh38Ensembl_1-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://May2017.archive.ensembl.org/Homo_sapiens/Gene/Summary?db=core;g=ENSG00000147889">GRCh38: Ensembl release 89: ENSG00000147889</a> – <a href="/wiki/Ensembl_genome_database_project" title="Ensembl genome database project">Ensembl</a>, May 2017</span> </li> <li id="cite_note-refGRCm38Ensembl-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-refGRCm38Ensembl_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-refGRCm38Ensembl_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-refGRCm38Ensembl_2-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://May2017.archive.ensembl.org/Mus_musculus/Gene/Summary?db=core;g=ENSMUSG00000044303">GRCm38: Ensembl release 89: ENSMUSG00000044303</a> – <a href="/wiki/Ensembl_genome_database_project" title="Ensembl genome database project">Ensembl</a>, May 2017</span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Biological+Chemistry&rft.atitle=The+human+tumor+suppressor+arf+interacts+with+spinophilin%2Fneurabin+II%2C+a+type+1+protein-phosphatase-binding+protein&rft.volume=276&rft.issue=17&rft.pages=14161-14169&rft.date=2001-04&rft_id=info%3Adoi%2F10.1074%2Fjbc.M006845200&rft_id=info%3Apmid%2F11278317&rft.aulast=Vivo&rft.aufirst=M&rft.au=Calogero%2C+RA&rft.au=Sansone%2C+F&rft.au=Calabr%C3%B2%2C+V&rft.au=Parisi%2C+T&rft.au=Borrelli%2C+L&rft.au=Saviozzi%2C+S&rft.au=La+Mantia%2C+G&rft_id=https%3A%2F%2Fdoi.org%2F10.1074%252Fjbc.M006845200&rfr_id=info%3Asid%2Fen.wikipedia.org%3AP16" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=P16&action=edit&section=20" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Genes%2C+p16">Genes,+p16</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgTracks?db=hg38&singleSearch=knownCanonical&position=CDKN2A"><i>CDKN2A</i></a> human gene location in the <a href="/wiki/UCSC_Genome_Browser" title="UCSC Genome Browser">UCSC Genome Browser</a>.</li> <li><a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgGene?db=hg38&hgg_type=knownGene&hgg_gene=CDKN2A"><i>CDKN2A</i></a> human gene details in the <a href="/wiki/UCSC_Genome_Browser" title="UCSC Genome Browser">UCSC Genome Browser</a>.</li></ul> <div class="navbox-styles"><style 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nochecker" style="line-height:130%"> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 210px;"><span typeof="mw:File"><a href="/wiki/File:PDB_1a5e_EBI.jpg" class="mw-file-description" title="1a5e: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 18 STRUCTURES"><img alt="1a5e: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 18 STRUCTURES" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bf/PDB_1a5e_EBI.jpg/180px-PDB_1a5e_EBI.jpg" decoding="async" width="180" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bf/PDB_1a5e_EBI.jpg/270px-PDB_1a5e_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bf/PDB_1a5e_EBI.jpg/360px-PDB_1a5e_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a></span></div> <div class="gallerytext"><b>1a5e</b>: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 18 STRUCTURES</div> </li> <li class="gallerybox" style="width: 215px"> <div 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<li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 210px;"><span typeof="mw:File"><a href="/wiki/File:PDB_1dc2_EBI.jpg" class="mw-file-description" title="1dc2: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 20 STRUCTURES"><img alt="1dc2: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 20 STRUCTURES" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/PDB_1dc2_EBI.jpg/180px-PDB_1dc2_EBI.jpg" decoding="async" width="180" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/PDB_1dc2_EBI.jpg/270px-PDB_1dc2_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c3/PDB_1dc2_EBI.jpg/360px-PDB_1dc2_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a></span></div> <div class="gallerytext"><b>1dc2</b>: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, 20 STRUCTURES</div> </li> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 210px;"><span typeof="mw:File"><a href="/wiki/File:PDB_2a5e_EBI.jpg" class="mw-file-description" title="2a5e: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, RESTRAINED MINIMIZED MEAN STRUCTURE"><img alt="2a5e: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, RESTRAINED MINIMIZED MEAN STRUCTURE" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/PDB_2a5e_EBI.jpg/180px-PDB_2a5e_EBI.jpg" decoding="async" width="180" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/PDB_2a5e_EBI.jpg/270px-PDB_2a5e_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a9/PDB_2a5e_EBI.jpg/360px-PDB_2a5e_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a></span></div> <div class="gallerytext"><b>2a5e</b>: SOLUTION NMR STRUCTURE OF TUMOR SUPPRESSOR P16INK4A, RESTRAINED MINIMIZED MEAN STRUCTURE</div> </li> </ul></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" 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href="/wiki/Cyclin_D1" title="Cyclin D1">D1</a>, <a href="/wiki/Cyclin_D2" title="Cyclin D2">D2</a>, <a href="/wiki/Cyclin_D3" title="Cyclin D3">D3</a>)</li> <li><a href="/wiki/Cyclin_E" title="Cyclin E">E</a> (<a href="/wiki/Cyclin_E1" title="Cyclin E1">E1</a>, <a href="/wiki/Cyclin_E2" title="Cyclin E2">E2</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">CDK</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cyclin-dependent_kinase_1" title="Cyclin-dependent kinase 1">1</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_2" title="Cyclin-dependent kinase 2">2</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_3" title="Cyclin-dependent kinase 3">3</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">4</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_5" title="Cyclin-dependent kinase 5">5</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_6" title="Cyclin-dependent kinase 6">6</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_7" title="Cyclin-dependent kinase 7">7</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_8" title="Cyclin-dependent kinase 8">8</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_9" title="Cyclin-dependent kinase 9">9</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_10" title="Cyclin-dependent kinase 10">10</a></li> <li><a href="/wiki/CDC2L2" title="CDC2L2">11A</a></li> <li><a href="/wiki/CDC2L1" title="CDC2L1">11B</a></li> <li><a href="/wiki/CDK12" title="CDK12">12</a></li> <li><a href="/wiki/CDK13" title="CDK13">13</a></li> <li><a href="/wiki/PFTK1" title="PFTK1">14</a></li> <li><a href="/wiki/CDK-activating_kinase" title="CDK-activating kinase">CDK-activating kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cyclin-dependent_kinase_inhibitor_protein" title="Cyclin-dependent kinase inhibitor protein">CDK inhibitor</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/INK4" title="INK4">INK4a/ARF</a> (<a href="/wiki/P14arf" title="P14arf">p14arf</a>/<a class="mw-selflink selflink">p16</a>, <a href="/wiki/CDKN2B" title="CDKN2B">p15</a>, <a href="/wiki/CDKN2C" title="CDKN2C">p18</a>, <a href="/wiki/CDKN2D" title="CDKN2D">p19</a>)</li> <li><a href="/wiki/Cell_cycle#Inhibitors" title="Cell cycle">cip/kip</a> (<a href="/wiki/P21" title="P21">p21</a>, <a href="/wiki/CDKN1B" title="CDKN1B">p27</a>, <a href="/wiki/Cyclin-dependent_kinase_inhibitor_1C" title="Cyclin-dependent kinase inhibitor 1C">p57</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/P53_p63_p73_family" title="P53 p63 p73 family">P53 p63 p73 family</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/P53" title="P53">p53</a></li> <li><a href="/wiki/TP63" title="TP63">p63</a></li> <li><a href="/wiki/P73" title="P73">p73</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cdk1" class="mw-redirect" title="Cdk1">Cdc2</a></li> <li><a href="/wiki/Cdc25" title="Cdc25">Cdc25</a></li> <li><a href="/wiki/CDC42" title="CDC42">Cdc42</a></li> <li><a href="/wiki/Cellular_apoptosis_susceptibility_protein" title="Cellular apoptosis susceptibility protein">Cellular apoptosis susceptibility protein</a></li> <li><a href="/wiki/E2F" title="E2F">E2F</a></li> <li><a href="/wiki/Maturation_promoting_factor" title="Maturation promoting factor">Maturation promoting factor</a></li> <li><a href="/wiki/Wee1" title="Wee1">Wee</a></li> <li><a href="/wiki/Cullin" title="Cullin">Cullin</a> (<a href="/wiki/CUL7" title="CUL7">CUL7</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Phases and<br />checkpoints</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Interphase" title="Interphase">Interphase</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub> phase</a></li> <li><a href="/wiki/S_phase" title="S phase">S phase</a></li> <li><a href="/wiki/G2_phase" title="G2 phase">G<sub>2</sub> phase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_division" title="Cell division">M phase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitosis" title="Mitosis">Mitosis</a> (<a href="/wiki/Preprophase" title="Preprophase">Preprophase</a></li> <li><a href="/wiki/Prophase" title="Prophase">Prophase</a></li> <li><a href="/wiki/Prometaphase" title="Prometaphase">Prometaphase</a></li> <li><a href="/wiki/Metaphase" title="Metaphase">Metaphase</a></li> <li><a href="/wiki/Anaphase" title="Anaphase">Anaphase</a></li> <li><a href="/wiki/Telophase" title="Telophase">Telophase</a>)</li> <li><a href="/wiki/Cytokinesis" title="Cytokinesis">Cytokinesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">Cell cycle checkpoints</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Restriction_point" title="Restriction point">Restriction point</a></li> <li><a href="/wiki/Spindle_checkpoint" title="Spindle checkpoint">Spindle checkpoint</a></li> <li><a href="/wiki/Postreplication_checkpoint" title="Postreplication checkpoint">Postreplication checkpoint</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other cellular phases</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apoptosis" title="Apoptosis">Apoptosis</a></li> <li><a href="/wiki/G0_phase" title="G0 phase">G<sub>0</sub> phase</a></li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" 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factor">Growth factors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Platelet-derived_growth_factor" title="Platelet-derived growth factor">c-Sis/PDGF</a></li> <li><a href="/wiki/Hepatocyte_growth_factor" title="Hepatocyte growth factor">HGF</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptor</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CDH1_(gene)" class="mw-redirect" title="CDH1 (gene)">CDH1</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/PTCH1" title="PTCH1">PTCH1</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/TGF_beta_receptor_2" title="TGF beta receptor 2">TGF beta receptor 2</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">Receptor tyrosine kinase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/ErbB" title="ErbB">ErbB</a>/<a href="/wiki/Epidermal_growth_factor_receptor" title="Epidermal growth factor receptor">c-ErbB</a> <ul><li><a href="/wiki/HER2/neu" class="mw-redirect" title="HER2/neu">HER2/neu</a></li> <li><a href="/wiki/ERBB3" title="ERBB3">Her 3</a></li></ul></li> <li><a href="/wiki/C-Met" class="mw-redirect" title="C-Met">c-Met</a></li> <li><a href="/wiki/RET_proto-oncogene" title="RET proto-oncogene">c-Ret</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CD117" class="mw-redirect" title="CD117">c-Kit</a></li> <li><a href="/wiki/CD135" title="CD135">Flt3</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular signaling P+Ps</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Beta-catenin" class="mw-redirect" title="Beta-catenin">Beta-catenin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adenomatosis_polyposis_coli" class="mw-redirect" title="Adenomatosis polyposis coli">APC</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mothers_against_decapentaplegic_homolog_2" title="Mothers against decapentaplegic homolog 2">SMAD2</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_4" title="Mothers against decapentaplegic homolog 4">SMAD4</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/AKT" class="mw-redirect" title="AKT">c-Akt</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/PTEN_(gene)" title="PTEN (gene)">PTEN</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Merlin_(protein)" title="Merlin (protein)">Neurofibromin 2/Merlin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ras_(protein)" class="mw-redirect" title="Ras (protein)">c-Ras</a></li> <li><a href="/wiki/HRAS" title="HRAS">HRAS</a></li> <li><a href="/wiki/C-Raf" title="C-Raf">c-Raf</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Neurofibromin" title="Neurofibromin">Neurofibromin 1</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/unknown</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Proto-oncogene_tyrosine-protein_kinase_Src" title="Proto-oncogene tyrosine-protein kinase Src">c-Src</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Maspin" title="Maspin">Maspin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_cycle" title="Cell cycle">Cell cycle</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4</a></li> <li><a href="/wiki/Cyclin_D" title="Cyclin D">Cyclin D</a></li> <li><a href="/wiki/Cyclin_E" title="Cyclin E">Cyclin E</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/P53" title="P53">p53</a></li> <li><a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">pRb</a></li> <li><a href="/wiki/WT1" class="mw-redirect" title="WT1">WT1</a></li> <li><a href="/wiki/P16_(gene)" class="mw-redirect" title="P16 (gene)">p16</a>/<a href="/wiki/P14arf" title="P14arf">p14arf</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a>/<a href="/wiki/Fanconi_anemia" title="Fanconi anemia">Fanconi</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/BRCA1" title="BRCA1">BRCA1</a></li> <li><a href="/wiki/BRCA2" title="BRCA2">BRCA2</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ubiquitin_ligase" title="Ubiquitin ligase">Ubiquitin ligase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CBL_(gene)" title="CBL (gene)">CBL</a></li> <li><a href="/wiki/Mdm2" title="Mdm2">MDM2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Von_Hippel%E2%80%93Lindau_tumor_suppressor" title="Von Hippel–Lindau tumor suppressor">VHL</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/AP-1_transcription_factor" title="AP-1 transcription factor">AP-1</a> <ul><li><a href="/wiki/C-Fos" class="mw-redirect" title="C-Fos">c-Fos</a></li> <li><a href="/wiki/C-jun" class="mw-redirect" title="C-jun">c-Jun</a></li></ul></li> <li><a href="/wiki/Myc" title="Myc">c-Myc</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/KLF6" title="KLF6">KLF6</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Apoptosis_inhibitor" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">Apoptosis inhibitor</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/SDHB" title="SDHB">SDHB</a></li> <li><a href="/wiki/SDHD" title="SDHD">SDHD</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/ungrouped</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bcl-2" title="Bcl-2">c-Bcl-2</a></li> <li><a href="/wiki/Notch_signaling" class="mw-redirect" title="Notch signaling">Notch</a></li> <li><a href="/wiki/Stathmin" title="Stathmin">Stathmin</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=P16&oldid=1229048989">https://en.wikipedia.org/w/index.php?title=P16&oldid=1229048989</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Genes_on_human_chromosome_9" title="Category:Genes on human chromosome 9">Genes on human chromosome 9</a></li><li><a href="/wiki/Category:Tumor_suppressor_genes" title="Category:Tumor 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