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Post-translational modification - Wikipedia

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class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle PTMs involving addition of functional groups subsection</span> </button> <ul id="toc-PTMs_involving_addition_of_functional_groups-sublist" class="vector-toc-list"> <li id="toc-Addition_by_an_enzyme_in_vivo" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Addition_by_an_enzyme_in_vivo"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Addition by an enzyme <i>in vivo</i></span> </div> </a> <ul id="toc-Addition_by_an_enzyme_in_vivo-sublist" class="vector-toc-list"> <li id="toc-Hydrophobic_groups_for_membrane_localization" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Hydrophobic_groups_for_membrane_localization"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.1</span> <span>Hydrophobic groups for membrane localization</span> </div> </a> <ul id="toc-Hydrophobic_groups_for_membrane_localization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cofactors_for_enhanced_enzymatic_activity" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Cofactors_for_enhanced_enzymatic_activity"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.2</span> <span>Cofactors for enhanced enzymatic activity</span> </div> </a> <ul id="toc-Cofactors_for_enhanced_enzymatic_activity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Modifications_of_translation_factors" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Modifications_of_translation_factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.3</span> <span>Modifications of translation factors</span> </div> </a> <ul id="toc-Modifications_of_translation_factors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Smaller_chemical_groups" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Smaller_chemical_groups"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.4</span> <span>Smaller chemical groups</span> </div> </a> <ul id="toc-Smaller_chemical_groups-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Non-enzymatic_modifications_in_vivo" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-enzymatic_modifications_in_vivo"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Non-enzymatic modifications <i>in vivo</i></span> </div> </a> <ul id="toc-Non-enzymatic_modifications_in_vivo-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-enzymatic_additions_in_vitro" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-enzymatic_additions_in_vitro"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Non-enzymatic additions <i>in vitro</i></span> </div> </a> <ul id="toc-Non-enzymatic_additions_in_vitro-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Conjugation_with_other_proteins_or_peptides" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Conjugation_with_other_proteins_or_peptides"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Conjugation with other proteins or peptides</span> </div> </a> <ul id="toc-Conjugation_with_other_proteins_or_peptides-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Chemical_modification_of_amino_acids" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Chemical_modification_of_amino_acids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Chemical modification of amino acids</span> </div> </a> <ul id="toc-Chemical_modification_of_amino_acids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structural_changes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Structural_changes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Structural changes</span> </div> </a> <ul id="toc-Structural_changes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Statistics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Statistics"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Statistics</span> </div> </a> <button aria-controls="toc-Statistics-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 Statistics subsection</span> </button> <ul id="toc-Statistics-sublist" class="vector-toc-list"> <li id="toc-Common_PTMs_by_frequency" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Common_PTMs_by_frequency"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Common PTMs by frequency</span> </div> </a> <ul id="toc-Common_PTMs_by_frequency-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Common_PTMs_by_residue" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Common_PTMs_by_residue"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Common PTMs by residue</span> </div> </a> <ul id="toc-Common_PTMs_by_residue-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Databases_and_tools" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Databases_and_tools"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Databases and tools</span> </div> </a> <button aria-controls="toc-Databases_and_tools-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 Databases and tools subsection</span> </button> <ul id="toc-Databases_and_tools-sublist" class="vector-toc-list"> <li id="toc-List_of_resources" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#List_of_resources"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>List of resources</span> </div> </a> <ul id="toc-List_of_resources-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tools" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tools"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Tools</span> </div> </a> <ul id="toc-Tools-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Case_examples" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Case_examples"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Case examples</span> </div> </a> <ul id="toc-Case_examples-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">8</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">9</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">10</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" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Post-translational modification</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. Available in 29 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-29" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">29 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D8%B9%D8%AF%D9%8A%D9%84_%D9%85%D8%A7_%D8%A8%D8%B9%D8%AF_%D8%A7%D9%84%D8%AA%D8%B1%D8%AC%D9%85%D8%A9" title="تعديل ما بعد الترجمة – Arabic" lang="ar" hreflang="ar" data-title="تعديل ما بعد الترجمة" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Posttranslacijske_modifikacije" title="Posttranslacijske modifikacije – Bosnian" lang="bs" hreflang="bs" data-title="Posttranslacijske modifikacije" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Modificaci%C3%B3_posttraduccional" title="Modificació posttraduccional – Catalan" lang="ca" hreflang="ca" data-title="Modificació posttraduccional" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Posttransla%C4%8Dn%C3%AD_modifikace" title="Posttranslační modifikace – Czech" lang="cs" hreflang="cs" data-title="Posttranslační modifikace" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Posttranslationel_modifikation" title="Posttranslationel modifikation – Danish" lang="da" hreflang="da" data-title="Posttranslationel modifikation" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Posttranslationale_Modifikation" title="Posttranslationale Modifikation – German" lang="de" hreflang="de" data-title="Posttranslationale Modifikation" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Translatsioonij%C3%A4rgne_modifikatsioon" title="Translatsioonijärgne modifikatsioon – Estonian" lang="et" hreflang="et" data-title="Translatsioonijärgne modifikatsioon" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Modificaci%C3%B3n_postraduccional" title="Modificación postraduccional – Spanish" lang="es" hreflang="es" data-title="Modificación postraduccional" 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%BE%DB%8C%D8%B1%D8%A7%DB%8C%D8%B4_%D9%BE%D8%B3%D8%A7%D8%AA%D8%B1%D8%AC%D9%85%D9%87%E2%80%8C%D8%A7%DB%8C" title="پیرایش پساترجمه‌ای – Persian" lang="fa" hreflang="fa" data-title="پیرایش پساترجمه‌ای" 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/Modification_post-traductionnelle" title="Modification post-traductionnelle – French" lang="fr" hreflang="fr" data-title="Modification post-traductionnelle" 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/Modificaci%C3%B3n_postraducional" title="Modificación postraducional – Galician" lang="gl" hreflang="gl" data-title="Modificación postraducional" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%B2%88%EC%97%AD_%ED%9B%84_%EB%B3%80%ED%98%95" title="번역 후 변형 – Korean" lang="ko" hreflang="ko" data-title="번역 후 변형" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%80%D5%A5%D5%BF%D5%BF%D6%80%D5%A1%D5%B6%D5%BD%D5%AC%D5%A1%D6%81%D5%AB%D5%B8%D5%B6_%D5%B1%D6%87%D5%A1%D6%83%D5%B8%D5%AD%D5%B8%D6%82%D5%A9%D5%B5%D5%B8%D6%82%D5%B6" title="Հետտրանսլացիոն ձևափոխություն – Armenian" lang="hy" hreflang="hy" data-title="Հետտրանսլացիոն ձևափոխություն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Modifikasi_pascatranslasi" title="Modifikasi pascatranslasi – Indonesian" lang="id" hreflang="id" data-title="Modifikasi pascatranslasi" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Modificazione_post_traduzionale" title="Modificazione post traduzionale – Italian" lang="it" hreflang="it" data-title="Modificazione post traduzionale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A2%D7%99%D7%91%D7%95%D7%93_%D7%97%D7%9C%D7%91%D7%95%D7%A0%D7%99%D7%9D_%D7%9C%D7%90%D7%97%D7%A8_%D7%94%D7%AA%D7%A8%D7%92%D7%95%D7%9D" title="עיבוד חלבונים לאחר התרגום – Hebrew" lang="he" hreflang="he" data-title="עיבוד חלבונים לאחר התרגום" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Transzl%C3%A1ci%C3%B3_ut%C3%A1ni_m%C3%B3dos%C3%ADt%C3%A1s" title="Transzláció utáni módosítás – Hungarian" lang="hu" hreflang="hu" data-title="Transzláció utáni módosítás" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Pengubahsuaian_pascaterjemahan" title="Pengubahsuaian pascaterjemahan – Malay" lang="ms" hreflang="ms" data-title="Pengubahsuaian pascaterjemahan" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Posttranslationele_modificatie" title="Posttranslationele modificatie – Dutch" lang="nl" hreflang="nl" data-title="Posttranslationele modificatie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%BF%BB%E8%A8%B3%E5%BE%8C%E4%BF%AE%E9%A3%BE" title="翻訳後修飾 – Japanese" lang="ja" hreflang="ja" data-title="翻訳後修飾" 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/Modyfikacja_potranslacyjna" title="Modyfikacja potranslacyjna – Polish" lang="pl" hreflang="pl" data-title="Modyfikacja potranslacyjna" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Modifica%C3%A7%C3%A3o_p%C3%B3s-traducional_de_prote%C3%ADnas" title="Modificação pós-traducional de proteínas – Portuguese" lang="pt" hreflang="pt" data-title="Modificação pós-traducional de proteínas" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Modificare_post-transla%C8%9Bional%C4%83" title="Modificare post-translațională – Romanian" lang="ro" hreflang="ro" data-title="Modificare post-translațională" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BE%D1%81%D1%82%D1%82%D1%80%D0%B0%D0%BD%D1%81%D0%BB%D1%8F%D1%86%D0%B8%D0%BE%D0%BD%D0%BD%D0%B0%D1%8F_%D0%BC%D0%BE%D0%B4%D0%B8%D1%84%D0%B8%D0%BA%D0%B0%D1%86%D0%B8%D1%8F" title="Посттрансляционная модификация – Russian" lang="ru" hreflang="ru" data-title="Посттрансляционная модификация" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Posttranslaciona_modifikacija" title="Posttranslaciona modifikacija – Serbian" lang="sr" hreflang="sr" data-title="Posttranslaciona modifikacija" 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/Posttranslaciona_modifikacija" title="Posttranslaciona modifikacija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Posttranslaciona modifikacija" 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-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/%C3%87evrim_sonras%C4%B1_de%C4%9Fi%C5%9Fim" title="Çevrim sonrası değişim – Turkish" lang="tr" hreflang="tr" data-title="Çevrim sonrası değişim" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9F%D0%BE%D1%81%D1%82%D1%82%D1%80%D0%B0%D0%BD%D1%81%D0%BB%D1%8F%D1%86%D1%96%D0%B9%D0%BD%D0%B0_%D0%BC%D0%BE%D0%B4%D0%B8%D1%84%D1%96%D0%BA%D0%B0%D1%86%D1%96%D1%8F" title="Посттрансляційна модифікація – Ukrainian" lang="uk" hreflang="uk" 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class="mw-redirect" title="Post translational modification">Post translational modification</a>)</span></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">Chemical changes in proteins following their translation from mRNA</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Insulin_path.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Insulin_path.svg/300px-Insulin_path.svg.png" decoding="async" width="300" height="424" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Insulin_path.svg/450px-Insulin_path.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Insulin_path.svg/600px-Insulin_path.svg.png 2x" data-file-width="1488" data-file-height="2105" /></a><figcaption>Post-translational modification of <a href="/wiki/Insulin" title="Insulin">insulin</a>. At the top, the ribosome translates a mRNA sequence into a protein, insulin, and passes the protein through the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>, where it is cut, folded, and held in shape by disulfide (-S-S-) bonds. Then the protein passes through the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">golgi apparatus</a>, where it is packaged into a vesicle. In the vesicle, more parts are cut off, and it turns into mature insulin.</figcaption></figure> <p>In <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>, <b>post-translational modification</b> (<b>PTM</b>) is the <a href="/wiki/Covalent" class="mw-redirect" title="Covalent">covalent</a> process of changing <a href="/wiki/Protein" title="Protein">proteins</a> following <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein biosynthesis</a>. PTMs may involve <a href="/wiki/Enzymes" class="mw-redirect" title="Enzymes">enzymes</a> or occur spontaneously. Proteins are created by <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a>, which <a href="/wiki/Translation_(biology)" title="Translation (biology)">translate</a> <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> into <a href="/wiki/Polypeptide_chain" class="mw-redirect" title="Polypeptide chain">polypeptide chains</a>, which may then change to form the mature protein product. PTMs are important components in cell <a href="/wiki/Signal_transduction" title="Signal transduction">signalling</a>, as for example when <a href="/wiki/Prohormone" title="Prohormone">prohormones</a> are converted to <a href="/wiki/Hormone" title="Hormone">hormones</a>. </p><p>Post-translational modifications can occur on the <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> <a href="/wiki/Side_chain" title="Side chain">side chains</a> or at the protein's <a href="/wiki/C-terminus" title="C-terminus">C-</a> or <a href="/wiki/N-terminus" title="N-terminus">N-</a> termini.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> They can expand the chemical set of the 22 <a href="/wiki/Proteinogenic_amino_acid" title="Proteinogenic amino acid">amino acids</a> by changing an existing <a href="/wiki/Functional_group" title="Functional group">functional group</a> or adding a new one such as phosphate. <a href="/wiki/Phosphorylation" title="Phosphorylation">Phosphorylation</a> is highly effective for controlling the enzyme activity and is the most common change after translation. <sup id="cite_ref-khoury_2-0" class="reference"><a href="#cite_note-khoury-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> Many <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> and <a href="/wiki/Prokaryotic" class="mw-redirect" title="Prokaryotic">prokaryotic</a> proteins also have <a href="/wiki/Carbohydrate" title="Carbohydrate">carbohydrate</a> molecules attached to them in a process called <a href="/wiki/Glycosylation" title="Glycosylation">glycosylation</a>, which can promote <a href="/wiki/Protein_folding" title="Protein folding">protein folding</a> and improve stability as well as serving regulatory functions. Attachment of <a href="/wiki/Lipid" title="Lipid">lipid</a> molecules, known as <a href="/wiki/Lipidation" class="mw-redirect" title="Lipidation">lipidation</a>, often targets a protein or part of a protein attached to the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a>. </p><p>Other forms of post-translational modification consist of cleaving <a href="/wiki/Peptide_bond" title="Peptide bond">peptide bonds</a>, as in processing a <a href="/wiki/Propeptide" class="mw-redirect" title="Propeptide">propeptide</a> to a mature form or removing the initiator <a href="/wiki/Methionine" title="Methionine">methionine</a> residue. The formation of <a href="/wiki/Disulfide_bond" class="mw-redirect" title="Disulfide bond">disulfide bonds</a> from <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> residues may also be referred to as a post-translational modification.<sup id="cite_ref-lodish_3-0" class="reference"><a href="#cite_note-lodish-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> For instance, the peptide <a href="/wiki/Hormone" title="Hormone">hormone</a> <a href="/wiki/Insulin" title="Insulin">insulin</a> is cut twice after disulfide bonds are formed, and a <a href="/wiki/Propeptide" class="mw-redirect" title="Propeptide">propeptide</a> is removed from the middle of the chain; the resulting protein consists of two polypeptide chains connected by disulfide bonds. </p><p>Some types of post-translational modification are consequences of <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a>. <a href="/wiki/Carbonylation" title="Carbonylation">Carbonylation</a> is one example that targets the modified protein for degradation and can result in the formation of protein aggregates.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> Specific amino acid modifications can be used as <a href="/wiki/Biomarker" title="Biomarker">biomarkers</a> indicating oxidative damage.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> </p><p>Sites that often undergo post-translational modification are those that have a functional group that can serve as a <a href="/wiki/Nucleophile" title="Nucleophile">nucleophile</a> in the reaction: the <a href="/wiki/Hydroxyl" class="mw-redirect" title="Hydroxyl">hydroxyl</a> groups of <a href="/wiki/Serine" title="Serine">serine</a>, <a href="/wiki/Threonine" title="Threonine">threonine</a>, and <a href="/wiki/Tyrosine" title="Tyrosine">tyrosine</a>; the <a href="/wiki/Amine" title="Amine">amine</a> forms of <a href="/wiki/Lysine" title="Lysine">lysine</a>, <a href="/wiki/Arginine" title="Arginine">arginine</a>, and <a href="/wiki/Histidine" title="Histidine">histidine</a>; the <a href="/wiki/Thiolate" class="mw-redirect" title="Thiolate">thiolate</a> <a href="/wiki/Anion" class="mw-redirect" title="Anion">anion</a> of <a href="/wiki/Cysteine" title="Cysteine">cysteine</a>; the <a href="/wiki/Carboxylate" title="Carboxylate">carboxylates</a> of <a href="/wiki/Aspartate" class="mw-redirect" title="Aspartate">aspartate</a> and <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>; and the N- and C-termini. In addition, although the <a href="/wiki/Amide" title="Amide">amide</a> of <a href="/wiki/Asparagine" title="Asparagine">asparagine</a> is a weak nucleophile, it can serve as an attachment point for <a href="/wiki/Glycan" title="Glycan">glycans</a>. Rarer modifications can occur at oxidized <a href="/wiki/Methionine" title="Methionine">methionines</a> and at some <a href="/wiki/Methylene_group" title="Methylene group">methylene groups</a> in side chains.<sup id="cite_ref-walsh_7-0" class="reference"><a href="#cite_note-walsh-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> </p><p>Post-translational modification of proteins can be experimentally detected by a variety of techniques, including <a href="/wiki/Mass_spectrometry" title="Mass spectrometry">mass spectrometry</a>, <a href="/wiki/Eastern_blotting" class="mw-redirect" title="Eastern blotting">Eastern blotting</a>, and <a href="/wiki/Western_blotting" class="mw-redirect" title="Western blotting">Western blotting</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="PTMs_involving_addition_of_functional_groups">PTMs involving addition of functional groups</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=1" title="Edit section: PTMs involving addition of functional groups"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Addition_by_an_enzyme_in_vivo">Addition by an enzyme <i>in vivo</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=2" title="Edit section: Addition by an enzyme in vivo"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Hydrophobic_groups_for_membrane_localization">Hydrophobic groups for membrane localization</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=3" title="Edit section: Hydrophobic groups for membrane localization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Myristoylation" title="Myristoylation">myristoylation</a> (a type of <a href="/wiki/Acylation" title="Acylation">acylation</a>), attachment of <a href="/wiki/Myristate" class="mw-redirect" title="Myristate">myristate</a>, a C<sub>14</sub> saturated acid</li> <li><a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> (a type of acylation), attachment of <a href="/wiki/Palmitate" class="mw-redirect" title="Palmitate">palmitate</a>, a C<sub>16</sub> saturated acid</li> <li><a href="/wiki/Isoprenylation" class="mw-redirect" title="Isoprenylation">isoprenylation</a> or <a href="/wiki/Prenylation" title="Prenylation">prenylation</a>, the addition of an <a href="/wiki/Isoprenoid" class="mw-redirect" title="Isoprenoid">isoprenoid</a> group (e.g. <a href="/wiki/Farnesol" title="Farnesol">farnesol</a> and <a href="/wiki/Geranylgeraniol" title="Geranylgeraniol">geranylgeraniol</a>) <ul><li><a href="/wiki/Farnesylation" class="mw-redirect" title="Farnesylation">farnesylation</a></li> <li><a href="/wiki/Geranylgeranylation" title="Geranylgeranylation">geranylgeranylation</a></li></ul></li> <li><a href="/wiki/Glypiation" title="Glypiation">glypiation</a>, <a href="/wiki/GPI_anchor" class="mw-redirect" title="GPI anchor">glycosylphosphatidylinositol (GPI) anchor</a> formation via an amide bond to C-terminal tail</li></ul> <div class="mw-heading mw-heading4"><h4 id="Cofactors_for_enhanced_enzymatic_activity">Cofactors for enhanced enzymatic activity</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=4" title="Edit section: Cofactors for enhanced enzymatic activity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Lipoylation" class="mw-redirect" title="Lipoylation">lipoylation</a> (a type of acylation), attachment of a <a href="/wiki/Lipoate" class="mw-redirect" title="Lipoate">lipoate</a> (C<sub>8</sub>) functional group</li> <li><a href="/wiki/Flavin_group" title="Flavin group">flavin</a> moiety (<a href="/wiki/Flavin_mononucleotide" title="Flavin mononucleotide">FMN</a> or <a href="/wiki/Flavin_adenine_dinucleotide" title="Flavin adenine dinucleotide">FAD</a>) may be covalently attached</li> <li><a href="/wiki/Heme_C" title="Heme C">heme C</a> attachment via <a href="/wiki/Thioether" class="mw-redirect" title="Thioether">thioether</a> bonds with <a href="/wiki/Cysteine" title="Cysteine">cysteines</a></li> <li><a href="/wiki/Phosphopantetheinylation" class="mw-redirect" title="Phosphopantetheinylation">phosphopantetheinylation</a>, the addition of a 4'-phosphopantetheinyl moiety from <a href="/wiki/Coenzyme_A" title="Coenzyme A">coenzyme A</a>, as in fatty acid, polyketide, non-ribosomal peptide and leucine biosynthesis</li> <li><a href="/wiki/Retinylidene" class="mw-redirect" title="Retinylidene">retinylidene</a> <a href="/wiki/Schiff_base" title="Schiff base">Schiff base</a> formation</li></ul> <div class="mw-heading mw-heading4"><h4 id="Modifications_of_translation_factors">Modifications of translation factors</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=5" title="Edit section: Modifications of translation factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Diphthamide" title="Diphthamide">diphthamide</a> formation (on a histidine found in <a href="/wiki/EEF2" title="EEF2">eEF2</a>)</li> <li><a href="/w/index.php?title=Ethanolamine_phosphoglycerol&amp;action=edit&amp;redlink=1" class="new" title="Ethanolamine phosphoglycerol (page does not exist)">ethanolamine phosphoglycerol</a> attachment (on glutamate found in <a href="/wiki/Eukaryotic_translation_elongation_factor_1_alpha_1" title="Eukaryotic translation elongation factor 1 alpha 1">eEF1α</a>)<sup id="cite_ref-pmid2569467_8-0" class="reference"><a href="#cite_note-pmid2569467-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Hypusine" title="Hypusine">hypusine</a> formation (on conserved lysine of <a href="/wiki/EIF5A" title="EIF5A">eIF5A</a> (eukaryotic) and <a href="/w/index.php?title=AIF5A&amp;action=edit&amp;redlink=1" class="new" title="AIF5A (page does not exist)">aIF5A</a> (archaeal))</li> <li><a href="/wiki/Beta-Lysine" class="mw-redirect" title="Beta-Lysine">beta-Lysine</a> addition on a conserved lysine of the <a href="/wiki/Elongation_factor_P" title="Elongation factor P">elongation factor P</a> (EFP) in most bacteria.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> EFP is a homolog to <a href="/wiki/EIF5A" title="EIF5A">eIF5A</a> (eukaryotic) and <a href="/w/index.php?title=AIF5A&amp;action=edit&amp;redlink=1" class="new" title="AIF5A (page does not exist)">aIF5A</a> (archaeal) (see above).</li></ul> <div class="mw-heading mw-heading4"><h4 id="Smaller_chemical_groups">Smaller chemical groups</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=6" title="Edit section: Smaller chemical groups"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Acylation" title="Acylation">acylation</a>, e.g. <i>O</i>-acylation (<a href="/wiki/Esters" class="mw-redirect" title="Esters">esters</a>), <i>N</i>-acylation (<a href="/wiki/Amides" class="mw-redirect" title="Amides">amides</a>), <i>S</i>-acylation (<a href="/wiki/Thioesters" class="mw-redirect" title="Thioesters">thioesters</a>) <ul><li><a href="/wiki/Acetylation" title="Acetylation">acetylation</a>, the addition of an <a href="/wiki/Acetyl" class="mw-redirect" title="Acetyl">acetyl</a> group, either at the <a href="/wiki/N-terminus" title="N-terminus">N-terminus</a> of the protein or at <a href="/wiki/Lysine" title="Lysine">lysine</a> residues.<sup id="cite_ref-pmid29405707_10-0" class="reference"><a href="#cite_note-pmid29405707-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> The reverse is called <a href="/wiki/Deacetylation" class="mw-redirect" title="Deacetylation">deacetylation</a>.</li> <li><a href="/wiki/Formylation" title="Formylation">formylation</a></li></ul></li> <li><a href="/wiki/Alkylation" title="Alkylation">alkylation</a>, the addition of an <a href="/wiki/Alkyl" class="mw-redirect" title="Alkyl">alkyl</a> group, e.g. <a href="/wiki/Methyl" class="mw-redirect" title="Methyl">methyl</a>, <a href="/wiki/Ethyl_group" title="Ethyl group">ethyl</a> <ul><li><a href="/wiki/Methylation" title="Methylation">methylation</a> the addition of a <a href="/wiki/Methyl" class="mw-redirect" title="Methyl">methyl</a> group, usually at <a href="/wiki/Lysine" title="Lysine">lysine</a> or <a href="/wiki/Arginine" title="Arginine">arginine</a> residues. The reverse is called <a href="/wiki/Demethylation" title="Demethylation">demethylation</a>.</li></ul></li> <li><span id="amidation">amidation</span> at C-terminus. Formed by oxidative dissociation of a C-terminal Gly residue.<sup id="cite_ref-amidation_11-0" class="reference"><a href="#cite_note-amidation-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Amide" title="Amide">amide</a> bond formation <ul><li><a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> addition <ul><li><a href="/wiki/Arginylation" title="Arginylation">arginylation</a>, a <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a>-mediation addition</li> <li><a href="/wiki/Polyglutamylation" title="Polyglutamylation">polyglutamylation</a>, covalent linkage of <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamic acid</a> residues to the N-terminus of tubulin and some other proteins.<sup id="cite_ref-pmid1967194_12-0" class="reference"><a href="#cite_note-pmid1967194-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> (See <a href="/w/index.php?title=Tubulin_polyglutamylase&amp;action=edit&amp;redlink=1" class="new" title="Tubulin polyglutamylase (page does not exist)">tubulin polyglutamylase</a>)</li> <li><a href="/wiki/Polyglycylation" title="Polyglycylation">polyglycylation</a>, covalent linkage of one to more than 40 <a href="/wiki/Glycine" title="Glycine">glycine</a> residues to the <a href="/wiki/Tubulin" title="Tubulin">tubulin</a> C-terminal tail</li></ul></li></ul></li> <li><a href="/w/index.php?title=Butyrylation&amp;action=edit&amp;redlink=1" class="new" title="Butyrylation (page does not exist)">butyrylation</a></li> <li><a href="/wiki/Gamma-carboxylation" class="mw-redirect" title="Gamma-carboxylation">gamma-carboxylation</a> dependent on <a href="/wiki/Vitamin_K" title="Vitamin K">Vitamin K</a><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Glycosylation" title="Glycosylation">glycosylation</a>, the addition of a <a href="/wiki/Glycosyl" title="Glycosyl">glycosyl</a> group to either <a href="/wiki/Arginine" title="Arginine">arginine</a>, <a href="/wiki/Asparagine" title="Asparagine">asparagine</a>, <a href="/wiki/Cysteine" title="Cysteine">cysteine</a>, <a href="/wiki/Hydroxylysine" title="Hydroxylysine">hydroxylysine</a>, <a href="/wiki/Serine" title="Serine">serine</a>, <a href="/wiki/Threonine" title="Threonine">threonine</a>, <a href="/wiki/Tyrosine" title="Tyrosine">tyrosine</a>, or <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> resulting in a <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoprotein</a>. Distinct from <a href="/wiki/Glycation" title="Glycation">glycation</a>, which is regarded as a nonenzymatic attachment of sugars. <ul><li><a href="/wiki/O-GlcNAc" title="O-GlcNAc"><i>O</i>-GlcNAc</a>, addition of <i>N</i>-acetylglucosamine to serine or threonine residues in a β-glycosidic linkage</li> <li>polysialylation, addition of <a href="/wiki/Polysialic_acid" title="Polysialic acid">polysialic acid</a>, PSA, to <a href="/wiki/NCAM" class="mw-redirect" title="NCAM">NCAM</a></li></ul></li> <li><a href="/w/index.php?title=Malonylation&amp;action=edit&amp;redlink=1" class="new" title="Malonylation (page does not exist)">malonylation</a></li> <li><a href="/wiki/Protein_hydroxylation" class="mw-redirect" title="Protein hydroxylation">hydroxylation</a>: addition of an oxygen atom to the side-chain of a Pro or Lys residue</li> <li><a href="/wiki/Iodination" class="mw-redirect" title="Iodination">iodination</a>: addition of an iodine atom to the aromatic ring of a tyrosine residue (e.g. in <a href="/wiki/Thyroglobulin" title="Thyroglobulin">thyroglobulin</a>)</li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">nucleotide addition</a> such as <a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li> <li><a href="/wiki/Phosphate_ester" class="mw-redirect" title="Phosphate ester">phosphate ester</a> (<i>O</i>-linked) or <a href="/wiki/Phosphoramidate" title="Phosphoramidate">phosphoramidate</a> (<i>N</i>-linked) formation <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a>, the addition of a <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> group, usually to <a href="/wiki/Serine" title="Serine">serine</a>, <a href="/wiki/Threonine" title="Threonine">threonine</a>, and <a href="/wiki/Tyrosine" title="Tyrosine">tyrosine</a> (<i>O</i>-linked), or <a href="/wiki/Histidine" title="Histidine">histidine</a> (<i>N</i>-linked)</li> <li><a href="/wiki/Adenylylation" title="Adenylylation">adenylylation</a>, the addition of an <a href="/wiki/Adenylyl" class="mw-redirect" title="Adenylyl">adenylyl</a> moiety, usually to <a href="/wiki/Tyrosine" title="Tyrosine">tyrosine</a> (<i>O</i>-linked), or <a href="/wiki/Histidine" title="Histidine">histidine</a> and <a href="/wiki/Lysine" title="Lysine">lysine</a> (<i>N</i>-linked)</li> <li>uridylylation, the addition of an uridylyl-group (i.e. <a href="/wiki/Uridine_monophosphate" title="Uridine monophosphate">uridine monophosphate</a>, UMP), usually to tyrosine</li></ul></li> <li><a href="/wiki/Propionylation" title="Propionylation">propionylation</a></li> <li><a href="/wiki/Pyroglutamate" class="mw-redirect" title="Pyroglutamate">pyroglutamate</a> formation</li> <li><a href="/wiki/S-glutathionylation" class="mw-redirect" title="S-glutathionylation"><i>S</i>-glutathionylation</a></li> <li><a href="/wiki/S-nitrosylation" class="mw-redirect" title="S-nitrosylation"><i>S</i>-nitrosylation</a></li> <li><i>S</i>-sulfenylation (<i>aka</i> <i>S</i>-sulphenylation), reversible covalent addition of one oxygen atom to the <a href="/wiki/Thiol" title="Thiol">thiol</a> group of a <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> residue<sup id="cite_ref-CysOx_14-0" class="reference"><a href="#cite_note-CysOx-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup></li> <li><i>S</i>-sulfinylation, normally irreversible covalent addition of two oxygen atoms to the <a href="/wiki/Thiol" title="Thiol">thiol</a> group of a <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> residue<sup id="cite_ref-CysOx_14-1" class="reference"><a href="#cite_note-CysOx-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup></li> <li><i>S</i>-sulfonylation, normally irreversible covalent addition of three oxygen atoms to the <a href="/wiki/Thiol" title="Thiol">thiol</a> group of a <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> residue, resulting in the formation of a <a href="/wiki/Cysteic_acid" title="Cysteic acid">cysteic acid</a> residue<sup id="cite_ref-CysOx_14-2" class="reference"><a href="#cite_note-CysOx-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Succinylation" title="Succinylation">succinylation</a> addition of a <a href="/wiki/Succinyl" class="mw-redirect" title="Succinyl">succinyl</a> group to <a href="/wiki/Lysine" title="Lysine">lysine</a></li> <li><a href="/wiki/Tyrosine_sulfation" title="Tyrosine sulfation">sulfation</a>, the addition of a sulfate group to a <a href="/wiki/Tyrosine" title="Tyrosine">tyrosine</a>.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Non-enzymatic_modifications_in_vivo">Non-enzymatic modifications <i>in vivo</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=7" title="Edit section: Non-enzymatic modifications in vivo"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Examples of non-enzymatic PTMs are glycation, glycoxidation, nitrosylation, oxidation, succination, and lipoxidation.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> </p> <ul><li><a href="/wiki/Glycation" title="Glycation">glycation</a>, the addition of a sugar molecule to a protein without the controlling action of an enzyme.</li> <li><a href="/wiki/Carbamylation" class="mw-redirect" title="Carbamylation">carbamylation</a> the addition of <a href="/wiki/Isocyanic_acid" title="Isocyanic acid">Isocyanic acid</a> to a protein's N-terminus or the side-chain of Lys.<sup id="cite_ref-pmid21768218_16-0" class="reference"><a href="#cite_note-pmid21768218-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Carbonylation" title="Carbonylation">carbonylation</a> the addition of carbon monoxide to other organic/inorganic compounds.</li> <li>spontaneous <a href="/wiki/Isopeptide_bond" title="Isopeptide bond">isopeptide bond</a> formation, as found in many surface proteins of <a href="/wiki/Gram-positive_bacteria" title="Gram-positive bacteria">Gram-positive bacteria</a>.<sup id="cite_ref-pmid21055949_17-0" class="reference"><a href="#cite_note-pmid21055949-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Non-enzymatic_additions_in_vitro">Non-enzymatic additions <i>in vitro</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=8" title="Edit section: Non-enzymatic additions in vitro"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Biotinylation" title="Biotinylation">biotinylation</a>: covalent attachment of a biotin moiety using a biotinylation reagent, typically for the purpose of labeling a protein.</li> <li>carbamylation: the addition of Isocyanic acid to a protein's N-terminus or the side-chain of Lys or Cys residues, typically resulting from exposure to urea solutions.<sup id="cite_ref-stark_1960_18-0" class="reference"><a href="#cite_note-stark_1960-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup></li> <li>oxidation: addition of one or more Oxygen atoms to a susceptible side-chain, principally of Met, Trp, His or Cys residues. Formation of <a href="/wiki/Disulfide" title="Disulfide">disulfide</a> bonds between Cys residues.</li> <li><a href="/wiki/Pegylation" class="mw-redirect" title="Pegylation">pegylation</a>: covalent attachment of <a href="/wiki/Polyethylene_glycol" title="Polyethylene glycol">polyethylene glycol</a> (PEG) using a pegylation reagent, typically to the N-terminus or the side-chains of Lys residues. Pegylation is used to improve the efficacy of protein pharmaceuticals.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Conjugation_with_other_proteins_or_peptides">Conjugation with other proteins or peptides</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=9" title="Edit section: Conjugation with other proteins or peptides"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitination</a>, the <a href="/wiki/Covalent" class="mw-redirect" title="Covalent">covalent</a> linkage to the protein ubiquitin.</li> <li><a href="/wiki/SUMOylation" class="mw-redirect" title="SUMOylation">SUMOylation</a>, the covalent linkage to the <a href="/wiki/SUMO_protein" title="SUMO protein">SUMO protein</a> (Small Ubiquitin-related MOdifier)<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Neddylation" title="Neddylation">neddylation</a>, the covalent linkage to the <a href="/wiki/NEDD8" title="NEDD8">Nedd protein</a></li> <li>ISGylation, the covalent linkage to the <a href="/wiki/ISG15" title="ISG15">ISG15</a> protein (Interferon-Stimulated Gene 15)<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Pupylation" class="mw-redirect" title="Pupylation">pupylation</a>, the covalent linkage to the <a href="/wiki/Prokaryotic_ubiquitin-like_protein" title="Prokaryotic ubiquitin-like protein">prokaryotic ubiquitin-like protein</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="Chemical_modification_of_amino_acids">Chemical modification of amino acids</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=10" title="Edit section: Chemical modification of amino acids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Citrullination" title="Citrullination">citrullination</a>, or <b>deimination</b>, the conversion of <a href="/wiki/Arginine" title="Arginine">arginine</a> to <a href="/wiki/Citrulline" title="Citrulline">citrulline</a><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Deamidation" title="Deamidation">deamidation</a>, the conversion of <a href="/wiki/Glutamine" title="Glutamine">glutamine</a> to <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamic acid</a> or <a href="/wiki/Asparagine" title="Asparagine">asparagine</a> to <a href="/wiki/Aspartic_acid" title="Aspartic acid">aspartic acid</a></li> <li><a href="/w/index.php?title=Eliminylation&amp;action=edit&amp;redlink=1" class="new" title="Eliminylation (page does not exist)">eliminylation</a>, the conversion to an <a href="/wiki/Alkene" title="Alkene">alkene</a> by <a href="/wiki/Elimination_reaction" title="Elimination reaction">beta-elimination</a> of <a href="/wiki/Phosphothreonine" class="mw-redirect" title="Phosphothreonine">phosphothreonine</a> and <a href="/wiki/Phosphoserine" title="Phosphoserine">phosphoserine</a>, or <a href="/wiki/Dehydration_reaction" title="Dehydration reaction">dehydration</a> of <a href="/wiki/Threonine" title="Threonine">threonine</a> and <a href="/wiki/Serine" title="Serine">serine</a><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup></li></ul> <div class="mw-heading mw-heading2"><h2 id="Structural_changes">Structural changes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=11" title="Edit section: Structural changes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Disulfide_bridge" class="mw-redirect" title="Disulfide bridge">disulfide bridges</a>, the covalent linkage of two <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> amino acids</li> <li><a href="/w/index.php?title=Lysine-cysteine_bridge&amp;action=edit&amp;redlink=1" class="new" title="Lysine-cysteine bridge (page does not exist)">lysine-cysteine bridges</a>, the covalent linkage of 1 lysine and 1 or 2 cystine residues via an oxygen atom (NOS and SONOS bridges)<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup></li> <li><a href="/wiki/Proteolytic_cleavage" class="mw-redirect" title="Proteolytic cleavage">proteolytic cleavage</a>, cleavage of a protein at a peptide bond</li> <li><a href="/wiki/Isoaspartate" title="Isoaspartate">isoaspartate</a> formation, via the cyclisation of asparagine or aspartic acid amino-acid residues</li> <li><a href="/wiki/Racemization" title="Racemization">racemization</a> <ul><li>of <a href="/wiki/Serine" title="Serine">serine</a> by <a href="/wiki/Protein-serine_epimerase" title="Protein-serine epimerase">protein-serine epimerase</a></li> <li>of <a href="/wiki/Alanine" title="Alanine">alanine</a> in <a href="/wiki/Dermorphin" title="Dermorphin">dermorphin</a>, a frog <a href="/wiki/Opioid_peptide" title="Opioid peptide">opioid peptide</a></li> <li>of <a href="/wiki/Methionine" title="Methionine">methionine</a> in <a href="/wiki/Deltorphin" title="Deltorphin">deltorphin</a>, also a frog opioid peptide</li></ul></li> <li><a href="/wiki/Protein_splicing" title="Protein splicing">protein splicing</a>, self-catalytic removal of <a href="/wiki/Intein" class="mw-redirect" title="Intein">inteins</a> analogous to mRNA processing</li></ul> <div class="mw-heading mw-heading2"><h2 id="Statistics">Statistics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=12" title="Edit section: Statistics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Common_PTMs_by_frequency">Common PTMs by frequency</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=13" title="Edit section: Common PTMs by frequency"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2011, statistics of each post-translational modification experimentally and putatively detected have been compiled using proteome-wide information from the Swiss-Prot database.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> The 10 most common experimentally found modifications were as follows:<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p> <table class="wikitable"> <tbody><tr> <th>Frequency </th> <th>Modification </th></tr> <tr> <td>58383 </td> <td><a href="/wiki/Phosphorylation" title="Phosphorylation">Phosphorylation</a> </td></tr> <tr> <td>6751 </td> <td><a href="/wiki/Acetylation" title="Acetylation">Acetylation</a> </td></tr> <tr> <td>5526 </td> <td><a href="/wiki/N-linked_glycosylation" title="N-linked glycosylation">N-linked glycosylation</a> </td></tr> <tr> <td>2844 </td> <td><a href="#amidation">Amidation</a> </td></tr> <tr> <td>1619 </td> <td><a href="/wiki/Hydroxylation" title="Hydroxylation">Hydroxylation</a> </td></tr> <tr> <td>1523 </td> <td><a href="/wiki/Methylation" title="Methylation">Methylation</a> </td></tr> <tr> <td>1133 </td> <td><a href="/wiki/O-linked_glycosylation" title="O-linked glycosylation">O-linked glycosylation</a> </td></tr> <tr> <td>878 </td> <td><a href="/wiki/Ubiquitin" title="Ubiquitin">Ubiquitylation</a> </td></tr> <tr> <td>826 </td> <td><a href="/wiki/Pyroglutamic_acid" title="Pyroglutamic acid">Pyrrolidone carboxylic acid</a> </td></tr> <tr> <td>504 </td> <td><a href="/wiki/Sulfation" title="Sulfation">Sulfation</a> </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Common_PTMs_by_residue">Common PTMs by residue</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=14" title="Edit section: Common PTMs by residue"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some common post-translational modifications to specific amino-acid residues are shown below. Modifications occur on the side-chain unless indicated otherwise. </p> <table class="wikitable sortable" style="text-align: left"> <tbody><tr> <th>Amino Acid</th> <th>Abbrev.</th> <th>Modification </th></tr> <tr> <th><a href="/wiki/Alanine" title="Alanine">Alanine</a> </th> <td>Ala or A </td> <td><a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus) </td></tr> <tr> <th><a href="/wiki/Arginine" title="Arginine">Arginine</a> </th> <td>Arg or R </td> <td>deimination to <a href="/wiki/Citrullination" title="Citrullination">citrulline</a>, <a href="/wiki/Protein_methylation" title="Protein methylation">methylation</a> </td></tr> <tr> <th><a href="/wiki/Asparagine" title="Asparagine">Asparagine</a> </th> <td>Asn or N </td> <td><a href="/wiki/Deamidation" title="Deamidation">deamidation</a> to Asp or iso(Asp), <a href="/wiki/N-linked_glycosylation" title="N-linked glycosylation">N-linked glycosylation</a>, <a href="/wiki/Isopeptide_bond" title="Isopeptide bond">spontaneous isopeptide bond formation</a> </td></tr> <tr> <th><a href="/wiki/Aspartic_acid" title="Aspartic acid">Aspartic acid</a> </th> <td>Asp or D </td> <td><a href="/wiki/Deamidation" title="Deamidation">isomerization</a> to isoaspartic acid, <a href="/wiki/Isopeptide_bond" title="Isopeptide bond">spontaneous isopeptide bond formation</a> </td></tr> <tr> <th><a href="/wiki/Cysteine" title="Cysteine">Cysteine</a> </th> <td>Cys or C </td> <td><a href="/wiki/Disulfide" title="Disulfide">disulfide</a>-bond formation, oxidation to sulfenic, sulfinic or sulfonic acid, <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a>, <a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus), <a href="/wiki/S-nitrosylation" class="mw-redirect" title="S-nitrosylation">S-nitrosylation</a> </td></tr> <tr> <th><a href="/wiki/Glutamine" title="Glutamine">Glutamine</a> </th> <td>Gln or Q </td> <td>cyclization to <a href="/wiki/Pyroglutamic_acid" title="Pyroglutamic acid">pyroglutamic acid</a> (N-terminus), deamidation to <a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> or <a href="/wiki/Isopeptide_bond" title="Isopeptide bond">isopeptide bond</a> formation to a <a href="/wiki/Lysine" title="Lysine">lysine</a> by a <a href="/wiki/Transglutaminase" title="Transglutaminase">transglutaminase</a> </td></tr> <tr> <th><a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> </th> <td>Glu or E </td> <td>cyclization to <a href="/wiki/Pyroglutamic_acid" title="Pyroglutamic acid">Pyroglutamic acid</a> (N-terminus), <a href="/wiki/Gamma-carboxylation" class="mw-redirect" title="Gamma-carboxylation">gamma-carboxylation</a> </td></tr> <tr> <th><a href="/wiki/Glycine" title="Glycine">Glycine</a> </th> <td>Gly or G </td> <td>N-<a href="/wiki/Myristoylation" title="Myristoylation">Myristoylation</a> (N-terminus), <a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus) </td></tr> <tr> <th><a href="/wiki/Histidine" title="Histidine">Histidine</a> </th> <td>His or H </td> <td><a href="/wiki/Histidine_kinase" title="Histidine kinase">Phosphorylation</a> </td></tr> <tr> <th><a href="/wiki/Isoleucine" title="Isoleucine">Isoleucine</a> </th> <td>Ile or I </td> <td> </td></tr> <tr> <th><a href="/wiki/Leucine" title="Leucine">Leucine</a> </th> <td>Leu or L </td> <td> </td></tr> <tr> <th><a href="/wiki/Lysine" title="Lysine">Lysine</a> </th> <td>Lys or K </td> <td><a href="/wiki/Acetylation" title="Acetylation">acetylation</a>, <a href="/wiki/Ubiquitylation" class="mw-redirect" title="Ubiquitylation">ubiquitylation</a>, <a href="/wiki/SUMO_protein" title="SUMO protein">SUMOylation</a>, <a href="/wiki/Protein_methylation" title="Protein methylation">methylation</a>, <a href="/wiki/Hydroxylation" title="Hydroxylation">hydroxylation</a> leading to <a href="/wiki/Allysine" title="Allysine">allysine</a>, <a href="/wiki/Isopeptide_bond" title="Isopeptide bond">spontaneous isopeptide bond formation</a> </td></tr> <tr> <th><a href="/wiki/Methionine" title="Methionine">Methionine</a> </th> <td>Met or M </td> <td><a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus), N-linked Ubiquitination, oxidation to sulfoxide or sulfone </td></tr> <tr> <th><a href="/wiki/Phenylalanine" title="Phenylalanine">Phenylalanine</a> </th> <td>Phe or F </td> <td> </td></tr> <tr> <th><a href="/wiki/Proline" title="Proline">Proline</a> </th> <td>Pro or P </td> <td><a href="/wiki/Hydroxylation" title="Hydroxylation">hydroxylation</a> </td></tr> <tr> <th><a href="/wiki/Serine" title="Serine">Serine</a> </th> <td>Ser or S </td> <td><a href="/wiki/Phosphorylation" title="Phosphorylation">Phosphorylation</a>, <a href="/wiki/O-linked_glycosylation" title="O-linked glycosylation">O-linked glycosylation</a>, <a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus) </td></tr> <tr> <th><a href="/wiki/Threonine" title="Threonine">Threonine</a> </th> <td>Thr or T </td> <td><a href="/wiki/Phosphorylation" title="Phosphorylation">Phosphorylation</a>, <a href="/wiki/O-linked_glycosylation" title="O-linked glycosylation">O-linked glycosylation</a>, <a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus) </td></tr> <tr> <th><a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</a> </th> <td>Trp or W </td> <td>mono- or di-oxidation, formation of <a href="/wiki/Kynurenine" title="Kynurenine">kynurenine</a>, <a href="/wiki/Tryptophan_tryptophylquinone" title="Tryptophan tryptophylquinone">tryptophan tryptophylquinone</a> </td></tr> <tr> <th><a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a> </th> <td>Tyr or Y </td> <td><a href="/wiki/Tyrosine_sulfation" title="Tyrosine sulfation">sulfation</a>, phosphorylation </td></tr> <tr> <th><a href="/wiki/Valine" title="Valine">Valine</a> </th> <td>Val or V </td> <td><a href="/wiki/Acetylation" title="Acetylation">N-acetylation</a> (N-terminus) </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Databases_and_tools">Databases and tools</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=15" title="Edit section: Databases and tools"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Image_for_Wiki_2.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/89/Image_for_Wiki_2.jpg/440px-Image_for_Wiki_2.jpg" decoding="async" width="440" height="295" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/8/89/Image_for_Wiki_2.jpg 1.5x" data-file-width="487" data-file-height="327" /></a><figcaption>Flowchart of the process and the data sources to predict PTMs.<sup id="cite_ref-ReferenceA_26-0" class="reference"><a href="#cite_note-ReferenceA-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> </figcaption></figure> <p>Protein sequences contain sequence motifs that are recognized by modifying enzymes, and which can be documented or predicted in PTM databases. With the large number of different modifications being discovered, there is a need to document this sort of information in databases. PTM information can be collected through experimental means or predicted from high-quality, manually curated data. Numerous databases have been created, often with a focus on certain taxonomic groups (e.g. human proteins) or other features. </p> <div class="mw-heading mw-heading3"><h3 id="List_of_resources">List of resources</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=16" title="Edit section: List of resources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://www.phosphosite.org/">PhosphoSitePlus</a><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> – A database of comprehensive information and tools for the study of mammalian protein post-translational modification</li> <li><a href="/w/index.php?title=ProteomeScout&amp;action=edit&amp;redlink=1" class="new" title="ProteomeScout (page does not exist)">ProteomeScout</a><sup id="cite_ref-ReferenceB_28-0" class="reference"><a href="#cite_note-ReferenceB-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> – A database of proteins and post-translational modifications experimentally</li> <li><a href="/wiki/Human_Protein_Reference_Database" title="Human Protein Reference Database">Human Protein Reference Database</a><sup id="cite_ref-ReferenceB_28-1" class="reference"><a href="#cite_note-ReferenceB-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> – A database for different modifications and understand different proteins, their class, and function/process related to disease causing proteins</li> <li><a href="/wiki/PROSITE" title="PROSITE">PROSITE</a><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">&#91;</span>29<span class="cite-bracket">&#93;</span></a></sup> – A database of Consensus patterns for many types of PTM's including sites</li> <li><a href="/wiki/Protein_Information_Resource" title="Protein Information Resource">RESID</a><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup> – A database consisting of a collection of annotations and structures for PTMs.</li> <li><a rel="nofollow" class="external text" href="https://research.bioinformatics.udel.edu/iptmnet/">iPTMnet</a> <sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">&#91;</span>31<span class="cite-bracket">&#93;</span></a></sup>– A database that integrates PTM information from several knowledgbases and text mining results.</li> <li><a href="/w/index.php?title=DbPTM&amp;action=edit&amp;redlink=1" class="new" title="DbPTM (page does not exist)">dbPTM</a><sup id="cite_ref-ReferenceA_26-1" class="reference"><a href="#cite_note-ReferenceA-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> – A database that shows different PTM's and information regarding their chemical components/structures and a frequency for amino acid modified site</li> <li><a rel="nofollow" class="external text" href="https://www.uniprot.org/help/post-translational_modification">Uniprot</a> has PTM information although that may be less comprehensive than in more specialized databases.<figure typeof="mw:File/Thumb"><a href="/wiki/File:Image_for_Wiki_1.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Image_for_Wiki_1.jpg/440px-Image_for_Wiki_1.jpg" decoding="async" width="440" height="367" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Image_for_Wiki_1.jpg/660px-Image_for_Wiki_1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Image_for_Wiki_1.jpg/880px-Image_for_Wiki_1.jpg 2x" data-file-width="1200" data-file-height="1002" /></a><figcaption>Effect of PTMs on protein function and physiological processes.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure></li> <li><a rel="nofollow" class="external text" href="https://www.oglcnac.mcw.edu/">The <i>O</i>-GlcNAc Database</a><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">&#91;</span>33<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">&#91;</span>34<span class="cite-bracket">&#93;</span></a></sup> - A curated database for protein O-GlcNAcylation and referencing more than 14 000 protein entries and 10 000 <i>O</i>-GlcNAc sites.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Tools">Tools</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=17" title="Edit section: Tools"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>List of software for visualization of proteins and their PTMs </p> <ul><li><a href="/wiki/PyMOL" title="PyMOL">PyMOL</a><sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">&#91;</span>35<span class="cite-bracket">&#93;</span></a></sup> – introduce a set of common PTM's into protein models</li> <li><a href="/w/index.php?title=AWESOME&amp;action=edit&amp;redlink=1" class="new" title="AWESOME (page does not exist)">AWESOME</a><sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">&#91;</span>36<span class="cite-bracket">&#93;</span></a></sup> – Interactive tool to see the role of single nucleotide polymorphisms to PTM's</li> <li><a href="/wiki/UCSF_Chimera" title="UCSF Chimera">Chimera</a><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">&#91;</span>37<span class="cite-bracket">&#93;</span></a></sup> – Interactive Database to visualize molecules</li></ul> <div class="mw-heading mw-heading2"><h2 id="Case_examples">Case examples</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Post-translational_modification&amp;action=edit&amp;section=18" title="Edit section: Case examples"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-More_citations_needed_section plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Post-translational_modification" title="Special:EditPage/Post-translational modification">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>&#32;in this section. Unsourced material may be challenged and removed.</span> <span class="date-container"><i>(<span class="date">January 2016</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <ul><li>Cleavage and formation of <a href="/wiki/Disulfide_bridge" class="mw-redirect" title="Disulfide bridge">disulfide bridges</a> during the production of <a href="/wiki/Insulin" title="Insulin">insulin</a></li> <li>PTM of <a href="/wiki/Histone" title="Histone">histones</a> as regulation of <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a>: <a href="/wiki/RNA_polymerase_control_by_chromatin_structure" class="mw-redirect" title="RNA polymerase control by chromatin structure">RNA polymerase control by chromatin structure</a></li> <li>PTM of <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a> as regulation of transcription</li> <li>Cleavage of polypeptide chains as crucial for lectin specificity<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">&#91;</span>38<span class="cite-bracket">&#93;</span></a></sup></li></ul> <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=Post-translational_modification&amp;action=edit&amp;section=19" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Protein_targeting" title="Protein targeting">Protein targeting</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational regulation</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=Post-translational_modification&amp;action=edit&amp;section=20" 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-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</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 .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFPrattVoetVoet2006" class="citation book cs1"><a href="/wiki/Charlotte_W._Pratt" title="Charlotte W. Pratt">Pratt, Charlotte W.</a>; <a href="/wiki/Judith_G._Voet" title="Judith G. Voet">Voet, Judith G.</a>; <a href="/wiki/Donald_Voet" title="Donald Voet">Voet, Donald</a> (2006). <a rel="nofollow" class="external text" href="https://archive.org/details/fundamentalsofbi00voet_0"><i>Fundamentals of Biochemistry: Life at the Molecular Level</i></a> (2nd&#160;ed.). Hoboken, NJ: Wiley. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/9780471214953" title="Special:BookSources/9780471214953"><bdi>9780471214953</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a>&#160;<a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1280801548">1280801548</a>. 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href="https://pubmed.ncbi.nlm.nih.gov/17623706">17623706</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Bioinformatics&amp;rft.atitle=structureViz%3A+linking+Cytoscape+and+UCSF+Chimera&amp;rft.volume=23&amp;rft.issue=17&amp;rft.pages=2345-7&amp;rft.date=2007-09&amp;rft_id=info%3Adoi%2F10.1093%2Fbioinformatics%2Fbtm329&amp;rft_id=info%3Apmid%2F17623706&amp;rft.aulast=Morris&amp;rft.aufirst=JH&amp;rft.au=Huang%2C+CC&amp;rft.au=Babbitt%2C+PC&amp;rft.au=Ferrin%2C+TE&amp;rft_id=https%3A%2F%2Fdoi.org%2F10.1093%252Fbioinformatics%252Fbtm329&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3APost-translational+modification" class="Z3988"></span></span> </li> <li id="cite_note-38"><span class="mw-cite-backlink"><b><a href="#cite_ref-38">^</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="http://www.proteopedia.org/wiki/index.php/1tp8">"1tp8 - Proteopedia, life in 3D"</a>. <i>www.proteopedia.org</i>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=unknown&amp;rft.jtitle=www.proteopedia.org&amp;rft.atitle=1tp8+-+Proteopedia%2C+life+in+3D&amp;rft_id=http%3A%2F%2Fwww.proteopedia.org%2Fwiki%2Findex.php%2F1tp8&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3APost-translational+modification" 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=Post-translational_modification&amp;action=edit&amp;section=21" 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://ftp.uniprot.org/pub/databases/uniprot/current_release/knowledgebase/complete/docs/ptmlist.txt">Controlled vocabulary of post-translational modifications</a> in <a href="/wiki/Uniprot" class="mw-redirect" title="Uniprot">Uniprot</a></li> <li><a rel="nofollow" class="external text" href="https://www.uniprot.org/docs/ptmlist">List of posttranslational modifications in ExPASy</a></li> <li><a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/dcbo.cgi?type=KW;po=9991">Browse SCOP domains by PTM</a> — from the <a href="/wiki/DcGO" title="DcGO">dcGO</a> database</li> <li><a rel="nofollow" class="external text" href="http://www.cytoskeleton.com/about-signal-seeker-ptm-detection">Overview and description of commonly used post-translational modification detection techniques</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist 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.navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Protein_topics" title="Template:Protein topics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Protein_topics" title="Template talk:Protein topics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Protein_topics" title="Special:EditPage/Template:Protein topics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Proteins" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein" title="Protein">Proteins</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Processes</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/Protein_biosynthesis" title="Protein biosynthesis">Protein biosynthesis</a></li> <li><a class="mw-selflink selflink">Post-translational modification</a></li> <li><a href="/wiki/Protein_folding" title="Protein folding">Protein folding</a></li> <li><a href="/wiki/Protein_targeting" title="Protein targeting">Protein targeting</a></li> <li><a href="/wiki/Proteome" title="Proteome">Proteome</a></li> <li><a href="/wiki/Protein_methods" title="Protein methods">Protein methods</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/Protein_structure" title="Protein structure">Protein structure</a></li> <li><a href="/wiki/Protein_domain" title="Protein domain">Protein structural domains</a></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</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-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/List_of_proteins" title="List of proteins">List of proteins</a></li> <li><a href="/wiki/Membrane_protein" title="Membrane protein">Membrane protein</a></li> <li><a href="/wiki/Globular_protein" title="Globular protein">Globular protein</a> <ul><li><a href="/wiki/Globulin" title="Globulin">Globulin</a></li> <li><a href="/wiki/Edestin" title="Edestin">Edestin</a></li> <li><a href="/wiki/Albumin" title="Albumin">Albumin</a></li></ul></li> <li><a href="/wiki/Scleroprotein" class="mw-redirect" title="Scleroprotein">Fibrous protein</a></li> <li><a href="/wiki/Chromoprotein" title="Chromoprotein">Chromoprotein</a></li> <li><a href="/wiki/Photoreceptor_protein" title="Photoreceptor protein">Photoreceptor protein</a></li> <li><a href="/wiki/Biliprotein" title="Biliprotein">Biliprotein</a> <ul><li><a href="/wiki/Phycobiliprotein" title="Phycobiliprotein">Phycobiliprotein</a></li> <li><a href="/wiki/Phytochrome" title="Phytochrome">Phytochrome</a></li> <li><a href="/wiki/Lipocalin" title="Lipocalin">Lipocalin</a></li></ul></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Protein_primary_structure_and_posttranslational_modifications" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Protein_primary_structure" title="Template:Protein primary structure"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Protein_primary_structure" title="Template talk:Protein primary structure"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Protein_primary_structure" title="Special:EditPage/Template:Protein primary structure"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Protein_primary_structure_and_posttranslational_modifications" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein_primary_structure" title="Protein primary structure">Protein primary structure</a> and <a href="/wiki/Posttranslational_modification" class="mw-redirect" title="Posttranslational modification">posttranslational modifications</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">General</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/Peptide_bond" title="Peptide bond">Peptide bond</a></li> <li><a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">Protein biosynthesis</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">Proteolysis</a></li> <li><a href="/wiki/Racemization" title="Racemization">Racemization</a></li> <li><a href="/w/index.php?title=N%E2%80%93O_acyl_shift&amp;action=edit&amp;redlink=1" class="new" title="N–O acyl shift (page does not exist)">N–O acyl shift</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/N_terminus" class="mw-redirect" title="N terminus">N terminus</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/Acetylation" title="Acetylation">Acetylation</a></li> <li><a href="/wiki/Carbamylation" class="mw-redirect" title="Carbamylation">Carbamylation</a></li> <li><a href="/wiki/Formylation" title="Formylation">Formylation</a></li> <li><a href="/wiki/Glycation" title="Glycation">Glycation</a></li> <li><a href="/wiki/Methylation" title="Methylation">Methylation</a></li> <li><a href="/wiki/Myristoylation" title="Myristoylation">Myristoylation</a> (Gly)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/C_terminus" class="mw-redirect" title="C terminus">C terminus</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/Amide" title="Amide">Amidation</a></li> <li><a href="/wiki/Glycophosphatidylinositol" class="mw-redirect" title="Glycophosphatidylinositol">Glycosyl phosphatidylinositol (GPI)</a></li> <li><a href="/w/index.php?title=O-methylation&amp;action=edit&amp;redlink=1" class="new" title="O-methylation (page does not exist)">O-methylation</a></li> <li><a href="/wiki/Detyrosination" title="Detyrosination">Detyrosination</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Single specific <a href="/wiki/Amino_acid" title="Amino acid">AAs</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/Serine" title="Serine">Serine</a>/<a href="/wiki/Threonine" title="Threonine">Threonine</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/Phosphorylation" title="Phosphorylation">Phosphorylation</a></li> <li><a href="/wiki/Dephosphorylation" title="Dephosphorylation">Dephosphorylation</a></li> <li><a href="/wiki/Glycosylation" title="Glycosylation">Glycosylation</a></li> <li><a href="/wiki/O-GlcNAc" title="O-GlcNAc"><i>O</i>-GlcNAc</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</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/Phosphorylation" title="Phosphorylation">Phosphorylation</a></li> <li><a href="/wiki/Dephosphorylation" title="Dephosphorylation">Dephosphorylation</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li> <li><a href="/wiki/Tyrosine_sulfation" title="Tyrosine sulfation">Sulfation</a></li> <li><a href="/wiki/Porphyrin" title="Porphyrin">Porphyrin ring linkage</a></li> <li><a href="/wiki/Adenylylation" title="Adenylylation">Adenylylation</a></li> <li><a href="/wiki/Flavin_group" title="Flavin group">Flavin linkage</a></li> <li><a href="/wiki/Topaquinone" title="Topaquinone">Topaquinone (TPQ) formation</a></li> <li><a href="/wiki/Detyrosination" title="Detyrosination">Detyrosination</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cysteine" title="Cysteine">Cysteine</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/Palmitoylation" title="Palmitoylation">Palmitoylation</a></li> <li><a href="/wiki/Prenylation" title="Prenylation">Prenylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Aspartate" class="mw-redirect" title="Aspartate">Aspartate</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/Deamidation" title="Deamidation">Succinimide formation</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">Glutamate</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/Carboxylation" title="Carboxylation">Carboxylation</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li> <li><a href="/wiki/Methylation" title="Methylation">Methylation</a></li> <li><a href="/wiki/Polyglutamylation" title="Polyglutamylation">Polyglutamylation</a></li> <li><a href="/wiki/Polyglycylation" title="Polyglycylation">Polyglycylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Asparagine" title="Asparagine">Asparagine</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/Deamidation" title="Deamidation">Deamidation</a></li> <li><a href="/wiki/Glycosylation" title="Glycosylation">Glycosylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glutamine" title="Glutamine">Glutamine</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/Transglutamination" class="mw-redirect" title="Transglutamination">Transglutamination</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lysine" title="Lysine">Lysine</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/Methylation" title="Methylation">Methylation</a></li> <li><a href="/wiki/Acetylation" title="Acetylation">Acetylation</a></li> <li><a href="/wiki/Acylation" title="Acylation">Acylation</a></li> <li><a href="/wiki/Adenylylation" title="Adenylylation">Adenylylation</a></li> <li><a href="/wiki/Hydroxylation" title="Hydroxylation">Hydroxylation</a></li> <li><a href="/wiki/Ubiquitination" class="mw-redirect" title="Ubiquitination">Ubiquitination</a></li> <li><a href="/wiki/SUMO_protein" title="SUMO protein">Sumoylation</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li> <li><a href="/wiki/Deamination" title="Deamination">Deamination</a></li> <li><a href="/wiki/Allysine" title="Allysine">Oxidative deamination to aldehyde</a></li> <li><a href="/wiki/O-glycosylation" class="mw-redirect" title="O-glycosylation"><i>O</i>-glycosylation</a></li> <li><a href="/wiki/Imine" title="Imine">Imine formation</a></li> <li><a href="/wiki/Glycation" title="Glycation">Glycation</a></li> <li><a href="/wiki/Carbamylation" class="mw-redirect" title="Carbamylation">Carbamylation</a></li> <li><a href="/wiki/Succinylation" title="Succinylation">Succinylation</a></li> <li><a href="/w/index.php?title=Lactylation&amp;action=edit&amp;redlink=1" class="new" title="Lactylation (page does not exist)">Lactylation</a></li> <li><a href="/wiki/Propionylation" title="Propionylation">Propionylation</a></li> <li><a href="/w/index.php?title=Butyrylation&amp;action=edit&amp;redlink=1" class="new" title="Butyrylation (page does not exist)">Butyrylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Arginine" title="Arginine">Arginine</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/Citrullination" title="Citrullination">Citrullination</a></li> <li><a href="/wiki/Methylation" title="Methylation">Methylation</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Proline" title="Proline">Proline</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/Hydroxylation" title="Hydroxylation">Hydroxylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Histidine" title="Histidine">Histidine</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/Diphthamide" title="Diphthamide">Diphthamide</a> formation</li> <li><a href="/wiki/Adenylylation" title="Adenylylation">Adenylylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</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/Glycosylation#C-mannosylation" title="Glycosylation">C-mannosylation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Crosslinks between two <a href="/wiki/Amino_acid" title="Amino acid">AAs</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/Cysteine" title="Cysteine">Cysteine</a>–<a href="/wiki/Cysteine" title="Cysteine">Cysteine</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/Disulfide_bond" class="mw-redirect" title="Disulfide bond">Disulfide bond</a></li> <li><a href="/wiki/ADP-ribosylation" title="ADP-ribosylation">ADP-ribosylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Methionine" title="Methionine">Methionine</a>–<a href="/wiki/Hydroxylysine" title="Hydroxylysine">Hydroxylysine</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/Sulfilimine_bond" class="mw-redirect" title="Sulfilimine bond">Sulfilimine bond</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lysine" title="Lysine">Lysine</a>–<a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</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="/w/index.php?title=Lysine_tyrosylquinone&amp;action=edit&amp;redlink=1" class="new" title="Lysine tyrosylquinone (page does not exist)">Lysine tyrosylquinone (LTQ) formation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</a>–<a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</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/Tryptophan_tryptophylquinone" title="Tryptophan tryptophylquinone">Tryptophan tryptophylquinone (TTQ) formation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Crosslinks between three <a href="/wiki/Amino_acid" title="Amino acid">AAs</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/Serine" title="Serine">Serine</a>–<a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a>–<a href="/wiki/Glycine" title="Glycine">Glycine</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/Green_Fluorescent_Protein#Autocatalytic_formation_of_the_chromophore_in_wtGFP" class="mw-redirect" title="Green Fluorescent Protein">p-Hydroxybenzylidene-imidazolinone (HBI) formation</a> (chromophore)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Histidine" title="Histidine">Histidine</a>–<a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a>–<a href="/wiki/Glycine" title="Glycine">Glycine</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/Kaede_(protein)" title="Kaede (protein)">4-(p-hydroxybenzylidene)-5-imidazolinone (HBI) formation</a> (chromophore)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Alanine" title="Alanine">Alanine</a>–<a href="/wiki/Serine" title="Serine">Serine</a>–<a href="/wiki/Glycine" title="Glycine">Glycine</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/Histidine_ammonia-lyase" title="Histidine ammonia-lyase">Methylidene-imidazolone (MIO) formation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Crosslinks between four <a href="/wiki/Amino_acid" title="Amino acid">AAs</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="Allysine–Allysine–Allysine–Lysine" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Allysine" title="Allysine">Allysine</a>–<a href="/wiki/Allysine" title="Allysine">Allysine</a>–<a href="/wiki/Allysine" title="Allysine">Allysine</a>–<a href="/wiki/Lysine" title="Lysine">Lysine</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/Desmosine" title="Desmosine">Desmosine</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" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Posttranslational_modification" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Posttranslational_modification" title="Template:Posttranslational modification"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Posttranslational_modification" title="Template talk:Posttranslational modification"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Posttranslational_modification" title="Special:EditPage/Template:Posttranslational modification"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Posttranslational_modification" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Posttranslational modification</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Chaperone_(protein)" title="Chaperone (protein)">Chaperones</a>/<br /><a href="/wiki/Protein_folding" title="Protein folding">protein folding</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%"><a href="/wiki/Heat_shock_protein" title="Heat shock protein">Heat shock proteins</a>/<br /><a href="/wiki/Chaperonin" title="Chaperonin">Chaperonins</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/GroES" title="GroES">Hsp10/GroES</a></li> <li><a href="/wiki/Hsp27" title="Hsp27">Hsp27</a></li> <li><a href="/wiki/Heat_shock_protein_47" title="Heat shock protein 47">Hsp47</a></li> <li><a href="/wiki/HSP60" class="mw-redirect" title="HSP60">HSP60</a>/<a href="/wiki/GroEL" title="GroEL">GroEL</a></li></ul> <ul><li><a href="/wiki/Hsp40" class="mw-redirect" title="Hsp40">Hsp40/DnaJ</a> <ul><li><a href="/wiki/DNAJA1" title="DNAJA1">A1</a></li> <li><a href="/wiki/DNAJA2" title="DNAJA2">A2</a></li> <li><a href="/wiki/DNAJA3" title="DNAJA3">A3</a></li> <li><a href="/wiki/DNAJB1" title="DNAJB1">B1</a></li> <li><a href="/wiki/DNAJB2" title="DNAJB2">B2</a></li> <li><a href="/wiki/DNAJB11" title="DNAJB11">B11</a></li> <li><a href="/wiki/DNAJB4" title="DNAJB4">B4</a></li> <li><a href="/wiki/DNAJB6" title="DNAJB6">B6</a></li> <li><a href="/wiki/DNAJB9" title="DNAJB9">B9</a></li> <li><a href="/wiki/DNAJC1" title="DNAJC1">C1</a></li> <li><a href="/wiki/DNAJC3" title="DNAJC3">C3</a></li> <li><a href="/wiki/DNAJC5" title="DNAJC5">C5</a></li> <li><a href="/wiki/DNAJC6" class="mw-redirect" title="DNAJC6">C6</a></li> <li><a href="/wiki/DNAJC7" title="DNAJC7">C7</a></li> <li><a href="/wiki/DNAJC10" title="DNAJC10">C10</a></li> <li><a href="/wiki/DNAJC11" title="DNAJC11">C11</a></li> <li><a href="/wiki/DNAJC13" title="DNAJC13">C13</a></li> <li><a href="/wiki/DNAJC14" title="DNAJC14">C14</a></li> <li><a href="/wiki/DNAJC19" title="DNAJC19">C19</a></li></ul></li></ul> <ul><li><a href="/wiki/Hsp70" title="Hsp70">Hsp70</a> <ul><li><a href="/wiki/HSPA1A" title="HSPA1A">1A</a></li> <li><a href="/wiki/HSPA1B" title="HSPA1B">1B</a></li> <li><a href="/wiki/HSPA1L" title="HSPA1L">1L</a></li> <li><a href="/wiki/HSPA2" title="HSPA2">2</a></li> <li><a href="/wiki/HSPA4" title="HSPA4">4</a></li> <li><a href="/wiki/HSPA4L" title="HSPA4L">4L</a></li> <li><a href="/wiki/Binding_immunoglobulin_protein" title="Binding immunoglobulin protein">5</a></li> <li><a href="/wiki/HSPA6" title="HSPA6">6</a></li> <li><a href="/wiki/HSPA7" title="HSPA7">7</a></li> <li><a href="/wiki/HSPA8" title="HSPA8">8</a></li> <li><a href="/wiki/HSPA9" title="HSPA9">9</a></li> <li><a href="/wiki/HSPA12A" title="HSPA12A">12A</a></li> <li><a href="/wiki/HSPA14" title="HSPA14">14</a></li></ul></li></ul> <ul><li><a href="/wiki/Hsp90" title="Hsp90">Hsp90</a> <ul><li><a href="/wiki/Heat_shock_protein_90kDa_alpha_(cytosolic),_member_A1" title="Heat shock protein 90kDa alpha (cytosolic), member A1">α<sub>1</sub></a></li> <li><a href="/wiki/HSP90AA2" title="HSP90AA2">α<sub>2</sub></a></li> <li><a href="/wiki/HSP90AB1" title="HSP90AB1">β</a></li> <li><a href="/wiki/HSP90B1" title="HSP90B1">ER</a></li> <li><a href="/wiki/TRAP1" title="TRAP1">TRAP1</a></li></ul></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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alpha_crystallin" class="mw-redirect" title="Alpha crystallin">Alpha crystallin</a></li> <li><a href="/wiki/Clusterin" title="Clusterin">Clusterin</a></li> <li><a href="/wiki/Survival_of_motor_neuron" title="Survival of motor neuron">Survival of motor neuron</a> <ul><li><a href="/wiki/SMN1" title="SMN1">SMN1</a></li> <li><a href="/wiki/SMN2" title="SMN2">SMN2</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Protein_targeting" title="Protein targeting">Protein targeting</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/Signal_peptide" title="Signal peptide">Signal peptide</a></li> <li><a href="/wiki/Target_peptide" title="Target peptide">Mitochondrial targeting signal</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ubiquitin" title="Ubiquitin">Ubiquitin</a><br />(ubiquitylation)</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>E1 <a href="/wiki/Ubiquitin-activating_enzyme" title="Ubiquitin-activating enzyme">Ubiquitin-activating enzyme</a> <ul><li><a href="/wiki/UBE1" class="mw-redirect" title="UBE1">UBA1</a></li> <li><a href="/wiki/UBA2" title="UBA2">UBA2</a></li> <li><a href="/wiki/UBE1C" title="UBE1C">UBA3</a></li> <li><a href="/wiki/UBE1DC1" class="mw-redirect" title="UBE1DC1">UBA5</a></li> <li><a href="/wiki/UBE1L2" title="UBE1L2">UBA6</a></li> <li><a href="/wiki/UBE1L" title="UBE1L">UBA7</a></li> <li><a href="/wiki/ATG7" title="ATG7">ATG7</a></li> <li><a href="/wiki/APPBP1" title="APPBP1">NAE1</a></li> <li><a href="/wiki/SAE1" title="SAE1">SAE1</a></li></ul></li></ul> <ul><li>E2 <a href="/wiki/Ubiquitin-conjugating_enzyme" title="Ubiquitin-conjugating enzyme">Ubiquitin-conjugating enzyme</a> <ul><li><a href="/wiki/UBE2A" title="UBE2A">A</a></li> <li><a href="/wiki/UBE2B" title="UBE2B">B</a></li> <li><a href="/wiki/UBE2C" title="UBE2C">C</a></li> <li><a href="/wiki/UBE2D1" title="UBE2D1">D1</a></li> <li><a href="/wiki/UBE2D2" title="UBE2D2">D2</a></li> <li><a href="/wiki/UBE2D3" title="UBE2D3">D3</a></li> <li><a href="/wiki/UBE2E1" title="UBE2E1">E1</a></li> <li><a href="/wiki/UBE2E2" title="UBE2E2">E2</a></li> <li><a href="/wiki/UBE2E3" title="UBE2E3">E3</a></li> <li><a href="/wiki/UBE2G1" title="UBE2G1">G1</a></li> <li><a href="/wiki/UBE2G2" title="UBE2G2">G2</a></li> <li><a href="/wiki/UBE2H" title="UBE2H">H</a></li> <li><a href="/wiki/UBE2I" title="UBE2I">I</a></li> <li><a href="/wiki/UBE2J1" title="UBE2J1">J1</a></li> <li><a href="/w/index.php?title=UBE2J2&amp;action=edit&amp;redlink=1" class="new" title="UBE2J2 (page does not exist)">J2</a></li> <li><a href="/w/index.php?title=UBE2K&amp;action=edit&amp;redlink=1" class="new" title="UBE2K (page does not exist)">K</a></li> <li><a href="/w/index.php?title=UBE2L1&amp;action=edit&amp;redlink=1" class="new" title="UBE2L1 (page does not exist)">L1</a></li> <li><a href="/w/index.php?title=UBE2L2&amp;action=edit&amp;redlink=1" class="new" title="UBE2L2 (page does not exist)">L2</a></li> <li><a href="/wiki/UBE2L3" title="UBE2L3">L3</a></li> <li><a href="/w/index.php?title=UBE2L4&amp;action=edit&amp;redlink=1" class="new" title="UBE2L4 (page does not exist)">L4</a></li> <li><a href="/wiki/UBE2L6" title="UBE2L6">L6</a></li> <li><a href="/wiki/UBE2M" title="UBE2M">M</a></li> <li><a href="/wiki/UBE2N" title="UBE2N">N</a></li> <li><a href="/wiki/UBE2O" title="UBE2O">O</a></li> <li><a href="/w/index.php?title=UBE2Q1&amp;action=edit&amp;redlink=1" class="new" title="UBE2Q1 (page does not exist)">Q1</a></li> <li><a href="/w/index.php?title=UBE2Q2&amp;action=edit&amp;redlink=1" class="new" title="UBE2Q2 (page does not exist)">Q2</a></li> <li><a href="/wiki/CDC34" title="CDC34">R1</a> (CDC34)</li> <li><a href="/wiki/UBE2R2" title="UBE2R2">R2</a></li> <li><a href="/w/index.php?title=UBE2S&amp;action=edit&amp;redlink=1" class="new" title="UBE2S (page does not exist)">S</a></li> <li><a href="/wiki/UBE2V1" title="UBE2V1">V1</a></li> <li><a href="/wiki/UBE2V2" title="UBE2V2">V2</a></li> <li><a href="/wiki/UBE2Z" title="UBE2Z">Z</a></li></ul></li></ul> <ul><li>E3 <a href="/wiki/Ubiquitin_ligase" title="Ubiquitin ligase">Ubiquitin ligase</a> <ul><li><a href="/wiki/Von_Hippel%E2%80%93Lindau_tumor_suppressor" title="Von Hippel–Lindau tumor suppressor">VHL</a></li> <li><a href="/wiki/Cullin" title="Cullin">Cullin</a></li> <li><a href="/wiki/CBL_(gene)" title="CBL (gene)">CBL</a></li> <li><a href="/wiki/Mdm2" title="Mdm2">MDM2</a></li> <li><a href="/wiki/FANCL" title="FANCL">FANCL</a></li> <li><a href="/wiki/UBR1" title="UBR1">UBR1</a></li></ul></li></ul> <ul><li><a href="/wiki/Deubiquitinating_enzyme" title="Deubiquitinating enzyme">Deubiquitinating enzyme</a>: <a href="/wiki/Ataxin_3" title="Ataxin 3">Ataxin 3</a></li> <li><a href="/wiki/USP6" title="USP6">USP6</a></li> <li><a href="/wiki/CYLD_(gene)" title="CYLD (gene)">CYLD</a></li></ul> <ul><li><a href="/wiki/ATG3" title="ATG3">ATG3</a></li> <li><a href="/wiki/BIRC6" title="BIRC6">BIRC6</a></li> <li><a href="/w/index.php?title=UFC1_(gene)&amp;action=edit&amp;redlink=1" class="new" title="UFC1 (gene) (page does not exist)">UFC1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ubiquitin-like_protein" title="Ubiquitin-like protein">Ubiquitin-like proteins</a><br />(UBL)</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%"><a href="/wiki/SUMO_protein" title="SUMO protein">SUMO protein</a><br />(SUMOylation)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>E1 SUMO-activating enzyme <ul><li><a href="/wiki/SAE1" title="SAE1">SAE1</a></li> <li><a href="/wiki/SAE2_(yeast)" title="SAE2 (yeast)">SAE2</a></li></ul></li></ul> <ul><li>E2 SUMO-conjugating enzyme <ul><li><a href="/wiki/UBC9" class="mw-redirect" title="UBC9">UBC9</a></li></ul></li></ul> <ul><li>E3 SUMO ligase <ul><li><a href="/wiki/PIAS1" title="PIAS1">PIAS1</a></li> <li><a href="/wiki/PIAS2" class="mw-redirect" title="PIAS2">PIAS2</a></li> <li><a href="/wiki/PIAS3" title="PIAS3">PIAS3</a></li> <li><a href="/wiki/PIAS4" title="PIAS4">PIAS4</a></li></ul></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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/ISG15" title="ISG15">ISG15</a></li> <li><a href="/wiki/URM1" title="URM1">URM1</a></li> <li><a href="/wiki/UFM1" title="UFM1">UFM1</a></li> <li><a href="/wiki/NEDD8" title="NEDD8">NEDD8</a> (<a href="/wiki/Neddylation" title="Neddylation">neddylation</a>)</li> <li><a href="/wiki/FAT10" class="mw-redirect" title="FAT10">FAT10</a></li> <li><a href="/wiki/ATG8" title="ATG8">ATG8</a></li> <li><a href="/wiki/ATG12" title="ATG12">ATG12</a></li> <li><a href="/w/index.php?title=FUB1&amp;action=edit&amp;redlink=1" class="new" title="FUB1 (page does not exist)">FUB1</a></li> <li><a href="/w/index.php?title=Membrane-anchored_UBL&amp;action=edit&amp;redlink=1" class="new" title="Membrane-anchored UBL (page does not exist)">MUB</a></li> <li><a href="/wiki/UBL5" title="UBL5">UBL5</a></li></ul> <ul><li><a href="/wiki/Prokaryotic_ubiquitin-like_protein" title="Prokaryotic ubiquitin-like protein">Prokaryotic ubiquitin-like protein</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" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Gene_expression" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction<br />to genetics</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/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a href="/wiki/DNA" title="DNA">DNA</a>&#160;→ <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a>&#160;→ <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</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:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA&#160;/&#32;hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</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/Translation_(biology)" title="Translation (biology)">Translation</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:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a class="mw-selflink selflink">Post-translational modification</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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</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/Epigenetics" title="Epigenetics">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</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/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a></li></ul> </div></td></tr></tbody></table></div> <style 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