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<span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Structure</span> </div> </a> <ul id="toc-Structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sequence_variants" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Sequence_variants"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Sequence variants</span> </div> </a> <ul id="toc-Sequence_variants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_translation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Alternative_translation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Alternative translation</span> </div> </a> <ul id="toc-Alternative_translation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Post-translational_modifications" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Post-translational_modifications"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Post-translational modifications</span> </div> </a> <button aria-controls="toc-Post-translational_modifications-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 Post-translational modifications subsection</span> </button> <ul id="toc-Post-translational_modifications-sublist" class="vector-toc-list"> <li id="toc-Methylation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Methylation"> <div 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">Histone H4</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. 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title="Histona H4 – Catalan" lang="ca" hreflang="ca" data-title="Histona H4" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Histon_H4" title="Histon H4 – German" lang="de" hreflang="de" data-title="Histon H4" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Histona_H4" title="Histona H4 – Spanish" lang="es" hreflang="es" data-title="Histona H4" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%87%DB%8C%D8%B3%D8%AA%D9%88%D9%86_%D8%A7%DA%86%DB%B4" title="هیستون اچ۴ – Persian" lang="fa" hreflang="fa" data-title="هیستون اچ۴" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Histone_H4" title="Histone H4 – French" lang="fr" hreflang="fr" data-title="Histone H4" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Histona_H4" title="Histona H4 – Galician" lang="gl" hreflang="gl" data-title="Histona H4" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Histon_H4" title="Histon H4 – Croatian" lang="hr" hreflang="hr" data-title="Histon H4" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Istone_H4" title="Istone H4 – Italian" lang="it" hreflang="it" data-title="Istone H4" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/H4_hiszton" title="H4 hiszton – Hungarian" lang="hu" hreflang="hu" data-title="H4 hiszton" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%92%E3%82%B9%E3%83%88%E3%83%B3H4" title="ヒストンH4 – Japanese" lang="ja" hreflang="ja" data-title="ヒストンH4" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Histon_H4" title="Histon H4 – Polish" lang="pl" hreflang="pl" data-title="Histon H4" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Histon_H4" title="Histon H4 – Serbian" lang="sr" hreflang="sr" data-title="Histon H4" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Histon_H4" title="Histon H4 – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Histon H4" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%93%D1%96%D1%81%D1%82%D0%BE%D0%BD_H4" title="Гістон H4 – Ukrainian" lang="uk" hreflang="uk" data-title="Гістон H4" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q24721644#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet 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.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">H4 histone, family 3</th></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">H4F3</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/National_Center_for_Biotechnology_Information#Gene" title="National Center for Biotechnology Information">NCBI gene</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/gene?cmd=retrieve&dopt=default&list_uids=3023&rn=1">3023</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/HUGO_Gene_Nomenclature_Committee" title="HUGO Gene Nomenclature Committee">HGNC</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.genenames.org/data/gene-symbol-report/#!/hgnc_id/HGNC:4780">4780</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Swiss-Prot" class="mw-redirect" title="Swiss-Prot">UniProt</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P62805">P62805</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Other data</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Locus_(genetics)" title="Locus (genetics)">Locus</a></th><td class="infobox-data" style="background-color: #eee"><a href="/wiki/Chromosome_3_(human)" class="mw-redirect" title="Chromosome 3 (human)">Chr. 3</a> <i><a rel="nofollow" class="external text" href="https://omim.org/search/?index=geneMap&search=3q13.13">q13.13</a></i></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">Search for</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;">Structures</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://swissmodel.expasy.org/repository/uniprot/P62805">Swiss-model</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;">Domains</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/interpro/protein/P62805">InterPro</a></td></tr></tbody></table></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Basic_units_of_chromatin_structure.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Basic_units_of_chromatin_structure.svg/220px-Basic_units_of_chromatin_structure.svg.png" decoding="async" width="220" height="495" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Basic_units_of_chromatin_structure.svg/330px-Basic_units_of_chromatin_structure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Basic_units_of_chromatin_structure.svg/440px-Basic_units_of_chromatin_structure.svg.png 2x" data-file-width="512" data-file-height="1152" /></a><figcaption>Basic units of chromatin structure</figcaption></figure> <p><b>Histone H4</b> is one of the five main <a href="/wiki/Histone" title="Histone">histone</a> <a href="/wiki/Protein" title="Protein">proteins</a> involved in the structure of <a href="/wiki/Chromatin" title="Chromatin">chromatin</a> in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells. Featuring a main globular domain and a long <a href="/wiki/N-terminus" title="N-terminus">N-terminal tail</a>, H4 is involved with the structure of the <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a> of the 'beads on a string' organization. Histone proteins are highly post-translationally modified. Covalently bonded modifications include <a href="/wiki/Acetylation" title="Acetylation">acetylation</a> and <a href="/wiki/Methylation" title="Methylation">methylation</a> of the N-terminal tails. These modifications may alter <a href="/wiki/Gene_expression" title="Gene expression">expression of genes</a> located on DNA associated with its parent histone octamer.<sup id="cite_ref-Bhasin_2006_1-0" class="reference"><a href="#cite_note-Bhasin_2006-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-HartlFreilfelder&Snyder_2-0" class="reference"><a href="#cite_note-HartlFreilfelder&Snyder-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Histone H4 is an important protein in the structure and function of chromatin, where its sequence variants and variable modification states are thought to play a role in the dynamic and long term regulation of genes. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Genetics">Genetics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=1" title="Edit section: Genetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Histone H4 is encoded in multiple genes at different loci including: <a href="/wiki/HIST1H4A" title="HIST1H4A">HIST1H4A</a>, <a href="/wiki/HIST1H4B" title="HIST1H4B">HIST1H4B</a>, <a href="/wiki/HIST1H4C" title="HIST1H4C">HIST1H4C</a>, <a href="/wiki/HIST1H4D" title="HIST1H4D">HIST1H4D</a>, <a href="/wiki/HIST1H4E" title="HIST1H4E">HIST1H4E</a>, <a href="/wiki/HIST1H4F" title="HIST1H4F">HIST1H4F</a>, <a href="/wiki/HIST1H4G" title="HIST1H4G">HIST1H4G</a>, <a href="/wiki/HIST1H4H" title="HIST1H4H">HIST1H4H</a>, <a href="/wiki/HIST1H4I" title="HIST1H4I">HIST1H4I</a>, <a href="/wiki/HIST1H4J" title="HIST1H4J">HIST1H4J</a>, <a href="/wiki/HIST1H4K" title="HIST1H4K">HIST1H4K</a>, <a href="/wiki/HIST1H4L" title="HIST1H4L">HIST1H4L</a>, <a href="/wiki/HIST2H4A" title="HIST2H4A">HIST2H4A</a>, <a href="/w/index.php?title=HIST2H4B&action=edit&redlink=1" class="new" title="HIST2H4B (page does not exist)">HIST2H4B</a>, <a href="/wiki/HIST4H4" title="HIST4H4">HIST4H4</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=2" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Histone proteins are among the most highly conserved eukaryotic proteins. For example, the amino acid sequence of histone H4 from a pea and cow differ at only 2 out of the 102 positions. This evolutionary conservation suggests that the functions of histone proteins involve nearly all of their amino acids so that any change is deleterious to the cell. Most changes in histone sequences are lethal; the few that are not lethal cause changes in the pattern of gene expression as well as other abnormalities.<sup id="cite_ref-Alberts2008_3-0" class="reference"><a href="#cite_note-Alberts2008-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=3" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Histone H4 is a 102 to 135 amino acid protein which shares a <a href="/wiki/Structural_motif" title="Structural motif">structural motif</a>, known as the <a href="/wiki/Histone_fold" title="Histone fold">histone fold</a>, formed from three a-helices connected by two loops. Histone proteins <a href="/wiki/Histone_H3" title="Histone H3">H3</a> and H4 bind to form a H3-H4 dimer, two of these H3-H4 dimers combine to form a <a href="/wiki/Tetramer" title="Tetramer">tetramer</a>. This tetramer further combines with two H2a-H2b dimers to form the compact <a href="/wiki/Histone_octamer" title="Histone octamer">Histone octamer</a> core.<sup id="cite_ref-Alberts2008_3-1" class="reference"><a href="#cite_note-Alberts2008-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Sequence_variants">Sequence variants</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=4" title="Edit section: Sequence variants"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Histone H4 is one of the slowest evolving proteins. There are H4 genes that are <a href="/wiki/Constitutively_expressed" class="mw-redirect" title="Constitutively expressed">constitutively expressed</a> throughout the cell cycle that encode for proteins that are identical in sequence to the major H4.<sup id="cite_ref-Kamakaka2005_4-0" class="reference"><a href="#cite_note-Kamakaka2005-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Variants in human histone H4 were only recently discovered and are very rare.<sup id="cite_ref-:0_5-0" class="reference"><a href="#cite_note-:0-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_6-0" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_7-0" class="reference"><a href="#cite_note-:2-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Pathogenic <i><a href="/wiki/De_novo_mutation" title="De novo mutation">de novo</a></i> missense variants have been identified in six H4 genes (<a href="/wiki/HIST1H4C" title="HIST1H4C">HIST1H4C</a>, <a href="/wiki/HIST1H4D" title="HIST1H4D">HIST1H4D</a>, <a href="/wiki/HIST1H4E" title="HIST1H4E">HIST1H4E</a>, <a href="/wiki/HIST1H4F" title="HIST1H4F">HIST1H4F</a>, <a href="/wiki/HIST1H4I" title="HIST1H4I">HIST1H4I</a>, <a href="/wiki/HIST1H4J" title="HIST1H4J">HIST1H4J</a>) in 33 individuals total, all presenting with <a href="/wiki/Neurodevelopmental_disorder" title="Neurodevelopmental disorder">neurodevelopmental</a> features of <a href="/wiki/Intellectual_disability" title="Intellectual disability">intellectual disability</a> and motor and/or gross developmental delay, but with variable non-neurological features. Ten amino acids were affected, six of which were found recurrently. These mutations were located within either the H4 core globular domain (involved in protein-protein interaction) or C-terminal tail (involved in post-translational modification).<sup id="cite_ref-:0_5-1" class="reference"><a href="#cite_note-:0-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_6-1" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_7-1" class="reference"><a href="#cite_note-:2-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Alternative_translation">Alternative translation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=5" title="Edit section: Alternative translation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The Osteogenic Growth Peptide (OGP) is a 14-aa peptide produced from alternative translation of histone H4 mRNA, sharing the <a href="/wiki/C-terminal" class="mw-redirect" title="C-terminal">C-terminal</a> sequence <code>ALKRQGRTLYGFGG</code> of histone H4. Translation is initiated at the 85th amino acid of the histone H4 mRNA, resulting in a 19-aa peptide (preOGP). This is converted into OGP through the cleavage of 5 amino-terminal residues.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> It is found in human and rat circulation as well as regenerating bone marrow. In blood serum it is bound to <a href="/wiki/Alpha-2-Macroglobulin" title="Alpha-2-Macroglobulin">α2M</a> along with two other binding proteins that are not clearly identified. A specific receptor has not been identified, but some signaling pathways involved in its bone-regenaration function has been elucidated.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Post-translational_modifications">Post-translational modifications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=6" title="Edit section: Post-translational modifications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Eukaryotic organisms can produce small amounts of specialized variant core histones that differ in amino acid sequence from the main ones. These variants with a variety of covalent modifications on the N-terminal can be added to histones making possible different chromatin structures that are required for DNA function in higher eukaryotes. Potential modifications include methylation (mono-, di-, or tri-methylation) or acetylation on the tails.<sup id="cite_ref-Alberts2008_3-2" class="reference"><a href="#cite_note-Alberts2008-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Methylation">Methylation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=7" title="Edit section: Methylation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Histone methylation occurs on arginine, lysine and histidine amino acids residues. Mono-, di- or tri-methylation has been discovered on histone H2A, H3 and H4.<sup id="cite_ref-Kim2009_10-0" class="reference"><a href="#cite_note-Kim2009-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Histone methylation has been associated with various cellular functions such as transcription, DNA replication, and DNA damage response including repair, heterochromatin formation, and somatic cell reprogramming. Among these biological functions, transcriptional repression and activation are the most studied.<sup id="cite_ref-Kim2009_10-1" class="reference"><a href="#cite_note-Kim2009-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Studies have shown that H4R3 methylation by PRMT1 (a histone methyltransferase) appears to be essential in vivo for the establishment or maintenance of a wide range of “active” chromatin modifications. Also methylation of histone H4 by PRMT1 was sufficient to permit subsequent acetylation on the N-terminal tail. However, acetylation of H4 inhibits its methylation by PRMT1.<sup id="cite_ref-Huang2005_11-0" class="reference"><a href="#cite_note-Huang2005-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Acetylation">Acetylation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=8" title="Edit section: Acetylation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Acetylation of histones is thought to relax condensed heterochromatin as the negative charge of acetyl groups can repel the DNA phosphate backbone charges, thus reducing the histone binding affinity for DNA. This hypothesis was validated by the discovery of the <a href="/wiki/Histone_acetyltransferase" title="Histone acetyltransferase">histone acetyltransferase</a> (HAT) activity of several transcriptional activator complexes.<sup id="cite_ref-Kim2009_10-2" class="reference"><a href="#cite_note-Kim2009-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Histone acetylation influences chromatin structure in several ways. First, it can provide a tag for the binding of proteins that contain areas which recognize the acetylated tails. Secondly, it can block the function of chromatin remodelers.<sup id="cite_ref-Taylor2013_12-0" class="reference"><a href="#cite_note-Taylor2013-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Thirdly, it neutralizes the positive charge on lysines.<sup id="cite_ref-Taylor2013_12-1" class="reference"><a href="#cite_note-Taylor2013-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Acetylation of histone H4 on lysine 16 (<a href="/wiki/H4K16ac" title="H4K16ac">H4K16ac</a>) is especially important for chromatin structure and function in a variety of eukaryotes and is catalyzed by specific histone lysine acetyltransferases (HATs). H4K16 is particularly interesting because this is the only acetylatable site of the H4 N-terminal tail, and can influence the formation of a compact higher-order chromatin structure.<sup id="cite_ref-Taylor2013_12-2" class="reference"><a href="#cite_note-Taylor2013-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Hypoacetylation of H4K16 appears to cause delayed recruitment of <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> proteins to sites of <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage</a> in a mouse model of the premature aging syndrome Hutchinson Gilford <a href="/wiki/Progeria" title="Progeria">progeria</a>.<sup id="cite_ref-pmid21746928_13-0" class="reference"><a href="#cite_note-pmid21746928-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> H4K16Ac also has roles in transcriptional activation and the maintenance of <a href="/wiki/Euchromatin" title="Euchromatin">euchromatin</a>.<sup id="cite_ref-Shrogren-Knaak2006_14-0" class="reference"><a href="#cite_note-Shrogren-Knaak2006-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Additional acetylations include K31ac and K79ac.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="List_of_H4_modifications">List of H4 modifications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=9" title="Edit section: List of H4 modifications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/w/index.php?title=H4S1p&action=edit&redlink=1" class="new" title="H4S1p (page does not exist)">H4S1p</a> </p><p><a href="/wiki/H4R3me2" title="H4R3me2">H4R3me2</a> </p><p><a href="/wiki/H4K5ac" title="H4K5ac">H4K5ac</a> </p><p><a href="/wiki/H4K8ac" title="H4K8ac">H4K8ac</a> </p><p><a href="/wiki/H4K12ac" title="H4K12ac">H4K12ac</a> </p><p><a href="/wiki/H4K16ac" title="H4K16ac">H4K16ac</a> </p><p><a href="/w/index.php?title=H4K16adp&action=edit&redlink=1" class="new" title="H4K16adp (page does not exist)">H4K16adp</a> </p><p><a href="/wiki/H4K20me" title="H4K20me">H4K20me</a> </p><p><a href="/w/index.php?title=H4K31ac&action=edit&redlink=1" class="new" title="H4K31ac (page does not exist)">H4K31ac</a> </p><p><a href="/w/index.php?title=H4S47o-p&action=edit&redlink=1" class="new" title="H4S47o-p (page does not exist)">H4S47o-p</a> </p><p><a href="/w/index.php?title=H4K79ac&action=edit&redlink=1" class="new" title="H4K79ac (page does not exist)">H4K79ac</a> </p><p><a href="/wiki/H4K91ac" title="H4K91ac">H4K91ac</a> </p><p><a href="/w/index.php?title=H4K91ub&action=edit&redlink=1" class="new" title="H4K91ub (page does not exist)">H4K91ub</a><sup id="cite_ref-poster_16-0" class="reference"><a href="#cite_note-poster-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <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=Histone_H4&action=edit&section=10" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Histone_code#Histone_H4" title="Histone code">Histone code#Histone H4</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=Histone_H4&action=edit&section=11" 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" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Bhasin_2006-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bhasin_2006_1-0">^</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 a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output 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(May 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4146406">"The first pilot project of the consortium for top-down proteomics: a status report"</a>. <i>Proteomics</i>. <b>14</b> (10): 1130–1140. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fpmic.201300438">10.1002/pmic.201300438</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4146406">4146406</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24644084">24644084</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proteomics&rft.atitle=The+first+pilot+project+of+the+consortium+for+top-down+proteomics%3A+a+status+report&rft.volume=14&rft.issue=10&rft.pages=1130-1140&rft.date=2014-05&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4146406%23id-name%3DPMC&rft_id=info%3Apmid%2F24644084&rft_id=info%3Adoi%2F10.1002%2Fpmic.201300438&rft.aulast=Dang&rft.aufirst=X&rft.au=Scotcher%2C+J&rft.au=Wu%2C+S&rft.au=Chu%2C+RK&rft.au=Toli%C4%87%2C+N&rft.au=Ntai%2C+I&rft.au=Thomas%2C+PM&rft.au=Fellers%2C+RT&rft.au=Early%2C+BP&rft.au=Zheng%2C+Y&rft.au=Durbin%2C+KR&rft.au=Leduc%2C+RD&rft.au=Wolff%2C+JJ&rft.au=Thompson%2C+CJ&rft.au=Pan%2C+J&rft.au=Han%2C+J&rft.au=Shaw%2C+JB&rft.au=Salisbury%2C+JP&rft.au=Easterling%2C+M&rft.au=Borchers%2C+CH&rft.au=Brodbelt%2C+JS&rft.au=Agar%2C+JN&rft.au=Pa%C5%A1a-Toli%C4%87%2C+L&rft.au=Kelleher%2C+NL&rft.au=Young%2C+NL&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4146406&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHistone+H4" class="Z3988"></span></span> </li> <li id="cite_note-poster-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-poster_16-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.abcam.com/pathways/epigenetic-modifications-poster">"Epigenetic modifications poster | Abcam"</a><span class="reference-accessdate">. Retrieved <span class="nowrap">25 November</span> 2019</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Epigenetic+modifications+poster+%26%23124%3B+Abcam&rft_id=https%3A%2F%2Fwww.abcam.com%2Fpathways%2Fepigenetic-modifications-poster&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHistone+H4" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="See_also_2">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Histone_H4&action=edit&section=12" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a></li> <li><a href="/wiki/Histone" title="Histone">histone</a></li> <li><a href="/wiki/Chromatin" title="Chromatin">chromatin</a></li> <li>Other <a href="/wiki/Histone" title="Histone">histone</a> <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a> involved in chromatin:</li></ul> <dl><dd><ul><li><a href="/wiki/Histone_H1" title="Histone H1">H1</a></li> <li><a href="/wiki/Histone_H2A" title="Histone H2A">H2A</a></li> <li><a href="/wiki/Histone_H2B" title="Histone H2B">H2B</a></li> <li><a href="/wiki/Histone_H3" title="Histone H3">H3</a></li></ul></dd></dl> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol 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a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Chromosome_genetics" title="Template:Chromosome genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Chromosome_genetics" title="Template talk:Chromosome genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Chromosome_genetics" title="Special:EditPage/Template:Chromosome genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cytogenetics:_chromosomes" style="font-size:114%;margin:0 4em"><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a>: <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basic<br />concepts</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/Karyotype" title="Karyotype">Karyotype</a></li> <li><a href="/wiki/Ploidy" title="Ploidy">Ploidy</a></li> <li><a href="/wiki/Genome" title="Genome">Genetic material</a>/<a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Chromatin" title="Chromatin">Chromatin</a> <ul><li><a href="/wiki/Euchromatin" title="Euchromatin">Euchromatin</a></li> <li><a href="/wiki/Heterochromatin" title="Heterochromatin">Heterochromatin</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></li> <li><a href="/wiki/Chromatid" title="Chromatid">Chromatid</a></li> <li><a href="/wiki/Nucleosome" title="Nucleosome">Nucleosome</a></li> <li><a href="/wiki/Nuclear_organization" title="Nuclear organization">Nuclear organization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</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/Autosome" title="Autosome">Autosome</a>/<a href="/wiki/Sex-determination_system" title="Sex-determination system">Sex chromosome</a> (or <a href="/wiki/Sex_chromosome" title="Sex chromosome">allosome</a> or heterosome)</li> <li><a href="/wiki/Microchromosome" title="Microchromosome">Macrochromosome/Microchromosome</a></li> <li><a href="/wiki/Circular_bacterial_chromosome" class="mw-redirect" title="Circular bacterial chromosome">Circular chromosome</a>/<a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear chromosome</a></li> <li><a href="/wiki/Aneuploidy" title="Aneuploidy">Extra chromosome</a> (or accessory chromosome)</li> <li><a href="/wiki/Small_supernumerary_marker_chromosome" title="Small supernumerary marker chromosome">Supernumerary chromosome</a></li> <li><a href="/wiki/B_chromosome" title="B chromosome">A chromosome/B chromosome</a></li> <li><a href="/wiki/Lampbrush_chromosome" title="Lampbrush chromosome">Lampbrush chromosome</a></li> <li><a href="/wiki/Polytene_chromosome" title="Polytene chromosome">Polytene chromosome</a></li> <li><a href="/wiki/Dinokaryon" title="Dinokaryon">Dinoflagellate chromosomes</a></li> <li><a href="/wiki/Homologous_chromosome" title="Homologous chromosome">Homologous chromosome</a></li> <li><a href="/wiki/Isochromosome" title="Isochromosome">Isochromosome</a></li> <li><a href="/wiki/Satellite_chromosome" title="Satellite chromosome">Satellite chromosome</a></li> <li><i>Centromere position</i> <ul><li><a href="/wiki/Centromere#Metacentric" title="Centromere">Metacentric</a></li> <li><a href="/wiki/Submetacentric" class="mw-redirect" title="Submetacentric">Submetacentric</a></li> <li><a href="/wiki/Telocentric" class="mw-redirect" title="Telocentric">Telocentric</a></li> <li><a href="/wiki/Acrocentric" class="mw-redirect" title="Acrocentric">Acrocentric</a></li> <li><a href="/wiki/Holocentric_chromosome" title="Holocentric chromosome">Holocentric</a></li></ul></li> <li><i>Centromere number</i> <ul><li><a href="/wiki/Acentric_fragment" title="Acentric fragment">Acentric</a></li> <li><a href="/wiki/Monocentric" class="mw-redirect" title="Monocentric">Monocentric</a></li> <li><a href="/wiki/Dicentric_chromosome" title="Dicentric chromosome">Dicentric</a></li> <li><a href="/wiki/Polycentric_chromosome" title="Polycentric chromosome">Polycentric</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Processes<br />and evolution</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></li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a></li> <li><a href="/wiki/Chromosome_abnormality#Structural_abnormalities" title="Chromosome abnormality">Structural alterations</a> <ul><li><a href="/wiki/Chromosomal_inversion" title="Chromosomal inversion">Chromosomal inversion</a></li> <li><a href="/wiki/Chromosomal_translocation" title="Chromosomal translocation">Chromosomal translocation</a></li></ul></li> <li><a href="/wiki/Chromosome_abnormality#Numerical_disorders" title="Chromosome abnormality">Numerical alterations</a> <ul><li><a href="/wiki/Aneuploidy" title="Aneuploidy">Aneuploidy</a></li> <li><a href="/wiki/Ploidy" title="Ploidy">Euploidy</a></li> <li><a href="/wiki/Polyploidy" title="Polyploidy">Polyploidy</a></li> <li><a href="/wiki/Paleopolyploidy" title="Paleopolyploidy">Paleopolyploidy</a></li></ul></li> <li><a href="/wiki/Polyploidization" class="mw-redirect" title="Polyploidization">Polyploidization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Structures</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/Telomere" title="Telomere">Telomere</a>: <a href="/wiki/Telomere-binding_protein" title="Telomere-binding protein">Telomere-binding protein</a> (<a href="/wiki/TINF2" title="TINF2">TINF2</a>)</li> <li><a href="/wiki/Protamine" title="Protamine">Protamine</a></li></ul> </div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Histone" title="Histone">Histone</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/Histone_H1" title="Histone H1">H1</a></li> <li><a href="/wiki/Histone_H2A" title="Histone H2A">H2A</a></li> <li><a href="/wiki/Histone_H2B" title="Histone H2B">H2B</a></li> <li><a href="/wiki/Histone_H3" title="Histone H3">H3</a></li> <li><a class="mw-selflink selflink">H4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Centromere" title="Centromere">Centromere</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/CENPA" title="CENPA">A</a></li> <li><a href="/wiki/Centromere_protein_B" title="Centromere protein B">B</a></li> <li><a href="/wiki/CENPC1" title="CENPC1">C1</a></li> <li><a href="/w/index.php?title=CENPC2&action=edit&redlink=1" class="new" title="CENPC2 (page does not exist)">C2</a></li> <li><a href="/wiki/Centromere_protein_E" title="Centromere protein E">E</a></li> <li><a href="/wiki/CENPF" title="CENPF">F</a></li> <li><a href="/wiki/CENPH" title="CENPH">H</a></li> <li><a href="/wiki/CENPI" title="CENPI">I</a></li> <li><a href="/wiki/CENPJ" title="CENPJ">J</a></li> <li><a href="/wiki/CENPK" title="CENPK">K</a></li> <li><a href="/wiki/CENPM" title="CENPM">M</a></li> <li><a href="/wiki/CENPN" title="CENPN">N</a></li> <li><a href="/wiki/CENPO" title="CENPO">O</a></li> <li><a href="/w/index.php?title=CENPP&action=edit&redlink=1" class="new" title="CENPP (page does not exist)">P</a></li> <li><a href="/w/index.php?title=CENPQ&action=edit&redlink=1" class="new" title="CENPQ (page does not exist)">Q</a></li> <li><a href="/wiki/CENPT" title="CENPT">T</a></li></ul> </div></td></tr></tbody></table><div> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</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/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a> <ul><li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li></ul></li> <li><a href="/wiki/List_of_organisms_by_chromosome_count" title="List of organisms by chromosome count">List of organisms by chromosome count</a></li> <li><a href="/wiki/List_of_sequenced_eukaryotic_genomes" title="List of sequenced eukaryotic genomes">List of sequenced genomes</a></li> <li><a href="/wiki/International_System_for_Human_Cytogenetic_Nomenclature" class="mw-redirect" title="International System for Human Cytogenetic Nomenclature">International System for Human Cytogenetic Nomenclature</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 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