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Transfer RNA - Wikipedia
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<span>Structure</span> </div> </a> <button aria-controls="toc-Structure-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 Structure subsection</span> </button> <ul id="toc-Structure-sublist" class="vector-toc-list"> <li id="toc-Anticodon" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anticodon"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Anticodon</span> </div> </a> <ul id="toc-Anticodon-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Nomenclature" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Nomenclature"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Nomenclature</span> </div> </a> <ul id="toc-Nomenclature-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Aminoacylation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Aminoacylation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Aminoacylation</span> </div> </a> <ul id="toc-Aminoacylation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Binding_to_ribosome" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Binding_to_ribosome"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Binding to ribosome</span> </div> </a> <ul id="toc-Binding_to_ribosome-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-tRNA_genes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#tRNA_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>tRNA genes</span> </div> </a> <button aria-controls="toc-tRNA_genes-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 tRNA genes subsection</span> </button> <ul id="toc-tRNA_genes-sublist" class="vector-toc-list"> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> <li id="toc-Hypothetical_origin" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Hypothetical_origin"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1.1</span> <span>Hypothetical origin</span> </div> </a> <ul id="toc-Hypothetical_origin-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-tRNA-derived_fragments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#tRNA-derived_fragments"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>tRNA-derived fragments</span> </div> </a> <ul id="toc-tRNA-derived_fragments-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Engineered_tRNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Engineered_tRNAs"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Engineered tRNAs</span> </div> </a> <ul id="toc-Engineered_tRNAs-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-tRNA_biogenesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#tRNA_biogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>tRNA biogenesis</span> </div> </a> <ul id="toc-tRNA_biogenesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clinical_relevance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_relevance"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Clinical relevance</span> </div> </a> <ul id="toc-Clinical_relevance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" 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">Transfer RNA</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 52 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-52" 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">52 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%AD%D9%85%D8%B6_%D9%86%D9%88%D9%88%D9%8A_%D8%B1%D9%8A%D8%A8%D9%88%D8%B2%D9%8A_%D9%86%D8%A7%D9%82%D9%84" title="حمض نووي ريبوزي ناقل – Arabic" lang="ar" hreflang="ar" data-title="حمض نووي ريبوزي ناقل" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%9F%E0%A7%8D%E0%A6%B0%E0%A6%BE%E0%A6%A8%E0%A7%8D%E0%A6%B8%E0%A6%AB%E0%A6%BE%E0%A6%B0_%E0%A6%86%E0%A6%B0%E0%A6%8F%E0%A6%A8%E0%A6%8F" title="ট্রান্সফার আরএনএ – Bangla" lang="bn" hreflang="bn" data-title="ট্রান্সফার আরএনএ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/Cho%C3%A1n-%C3%AE_RNA" title="Choán-î RNA – Minnan" lang="nan" hreflang="nan" data-title="Choán-î RNA" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A2%D1%80%D0%B0%D0%BD%D1%81%D0%BF%D0%BE%D1%80%D1%82%D0%BD%D0%B0_%D0%A0%D0%9D%D0%9A" title="Транспортна РНК – Bulgarian" lang="bg" hreflang="bg" data-title="Транспортна РНК" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Transportna_RNK" title="Transportna RNK – Bosnian" lang="bs" hreflang="bs" data-title="Transportna RNK" 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/ARN_de_transfer%C3%A8ncia" title="ARN de transferència – Catalan" lang="ca" hreflang="ca" data-title="ARN de transferència" 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/TRNA" title="TRNA – Czech" lang="cs" hreflang="cs" data-title="TRNA" 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/TRNA" title="TRNA – Danish" lang="da" hreflang="da" data-title="TRNA" 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/TRNA" title="TRNA – German" lang="de" hreflang="de" data-title="TRNA" 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/Transpordi-RNA" title="Transpordi-RNA – Estonian" lang="et" hreflang="et" data-title="Transpordi-RNA" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9C%CE%B5%CF%84%CE%B1%CF%86%CE%BF%CF%81%CE%B9%CE%BA%CF%8C_RNA" title="Μεταφορικό RNA – Greek" lang="el" hreflang="el" data-title="Μεταφορικό RNA" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/ARN_de_transferencia" title="ARN de transferencia – Spanish" lang="es" hreflang="es" data-title="ARN de transferencia" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Transferentziako_RNA" title="Transferentziako RNA – Basque" lang="eu" hreflang="eu" data-title="Transferentziako RNA" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A2%D8%B1%D8%A7%D9%86%E2%80%8C%D8%A7%DB%8C_%D8%AD%D8%A7%D9%85%D9%84" title="آرانای حامل – Persian" lang="fa" hreflang="fa" data-title="آرانای حامل" 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/Acide_ribonucl%C3%A9ique_de_transfert" title="Acide ribonucléique de transfert – French" lang="fr" hreflang="fr" data-title="Acide ribonucléique de transfert" 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/ARN_transferente" title="ARN transferente – Galician" lang="gl" hreflang="gl" data-title="ARN transferente" 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/%EC%9A%B4%EB%B0%98_RNA" title="운반 RNA – Korean" lang="ko" hreflang="ko" data-title="운반 RNA" 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%93%D5%B8%D5%AD%D5%A1%D5%A4%D6%80%D5%B8%D5%B2_%D5%8C%D5%86%D4%B9" title="Փոխադրող ՌՆԹ – Armenian" lang="hy" hreflang="hy" data-title="Փոխադրող ՌՆԹ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%85%E0%A4%A8%E0%A5%8D%E0%A4%A4%E0%A4%B0%E0%A4%A3_%E0%A4%86%E0%A4%B0%E0%A4%8F%E0%A4%A8%E0%A4%8F" title="अन्तरण आरएनए – Hindi" lang="hi" hreflang="hi" data-title="अन्तरण आरएनए" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Prijenosni_RNK" title="Prijenosni RNK – Croatian" lang="hr" hreflang="hr" data-title="Prijenosni RNK" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/RNA_transfer" title="RNA transfer – Indonesian" lang="id" hreflang="id" data-title="RNA transfer" 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-is mw-list-item"><a href="https://is.wikipedia.org/wiki/TRNA" title="TRNA – Icelandic" lang="is" hreflang="is" data-title="TRNA" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/RNA_transfer" title="RNA transfer – Italian" lang="it" hreflang="it" data-title="RNA transfer" 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/TRNA" title="TRNA – Hebrew" lang="he" hreflang="he" data-title="TRNA" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A1%E1%83%90%E1%83%A2%E1%83%A0%E1%83%90%E1%83%9C%E1%83%A1%E1%83%9E%E1%83%9D%E1%83%A0%E1%83%A2%E1%83%9D_%E1%83%A0%E1%83%98%E1%83%91%E1%83%9D%E1%83%9C%E1%83%A3%E1%83%99%E1%83%9A%E1%83%94%E1%83%98%E1%83%9C%E1%83%98%E1%83%A1_%E1%83%9B%E1%83%9F%E1%83%90%E1%83%95%E1%83%90" title="სატრანსპორტო რიბონუკლეინის მჟავა – Georgian" lang="ka" hreflang="ka" data-title="სატრანსპორტო რიბონუკლეინის მჟავა" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A2%D0%B0%D1%81%D1%8B%D0%BC%D0%B0%D0%BB%D0%B4%D0%B0%D1%83%D1%88%D1%8B_%D0%A0%D0%9D%D2%9A" title="Тасымалдаушы РНҚ – Kazakh" lang="kk" hreflang="kk" data-title="Тасымалдаушы РНҚ" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%A2%D0%B0%D1%88%D1%83%D1%83%D1%87%D1%83_%D0%A0%D0%9D%D0%9A" title="Ташуучу РНК – Kyrgyz" lang="ky" hreflang="ky" data-title="Ташуучу РНК" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/TRNA" title="TRNA – Latin" lang="la" hreflang="la" data-title="TRNA" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/TRNR" title="TRNR – Lithuanian" lang="lt" hreflang="lt" data-title="TRNR" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Transzfer_RNS" title="Transzfer RNS – Hungarian" lang="hu" hreflang="hu" data-title="Transzfer RNS" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%A2%D1%80%D0%B0%D0%BD%D1%81%D0%BF%D0%BE%D1%80%D1%82%D0%BD%D0%B0_%D0%A0%D0%9D%D0%9A" title="Транспортна РНК – Macedonian" lang="mk" hreflang="mk" data-title="Транспортна РНК" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/RNA_pemindah" title="RNA pemindah – Malay" lang="ms" hreflang="ms" data-title="RNA pemindah" 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/Transfer-RNA" title="Transfer-RNA – Dutch" lang="nl" hreflang="nl" data-title="Transfer-RNA" 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/%E8%BB%A2%E7%A7%BBRNA" title="転移RNA – Japanese" lang="ja" hreflang="ja" data-title="転移RNA" 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-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/TRNA" title="TRNA – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="TRNA" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Acid_ribonucle%C3%AFc_de_transferiment" title="Acid ribonucleïc de transferiment – Occitan" lang="oc" hreflang="oc" data-title="Acid ribonucleïc de transferiment" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/TRNA" title="TRNA – Polish" lang="pl" hreflang="pl" data-title="TRNA" 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/ARN_transportador" title="ARN transportador – Portuguese" lang="pt" hreflang="pt" data-title="ARN transportador" 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/Acid_ribonucleic_de_transfer" title="Acid ribonucleic de transfer – Romanian" lang="ro" hreflang="ro" data-title="Acid ribonucleic de transfer" 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%A2%D0%A0%D0%9D%D0%9A" title="ТРНК – Russian" lang="ru" hreflang="ru" data-title="ТРНК" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Transferov%C3%A1_ribonukleov%C3%A1_kyselina" title="Transferová ribonukleová kyselina – Slovak" lang="sk" hreflang="sk" data-title="Transferová ribonukleová kyselina" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%A6%D8%A7%DA%95_%D8%A6%DB%8E%D9%86_%D8%A6%DB%95%DB%8C_%DA%AF%D9%88%DB%8E%D8%B2%DB%95%D8%B1%DB%95%D9%88%DB%95" title="ئاڕ ئێن ئەی گوێزەرەوە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="ئاڕ ئێن ئەی گوێزەرەوە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A2%D1%80%D0%B0%D0%BD%D1%81%D0%BF%D0%BE%D1%80%D1%82%D0%BD%D0%B0_%D0%A0%D0%9D%D0%9A" title="Транспортна РНК – Serbian" lang="sr" hreflang="sr" data-title="Транспортна РНК" 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/Transportna_RNK" title="Transportna RNK – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Transportna RNK" 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-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Siirt%C3%A4j%C3%A4-RNA" title="Siirtäjä-RNA – Finnish" lang="fi" hreflang="fi" data-title="Siirtäjä-RNA" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Transfer-RNA" title="Transfer-RNA – Swedish" lang="sv" hreflang="sv" data-title="Transfer-RNA" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%97%E0%B8%B5%E0%B8%AD%E0%B8%B2%E0%B8%A3%E0%B9%8C%E0%B9%80%E0%B8%AD%E0%B9%87%E0%B8%99%E0%B9%80%E0%B8%AD" title="ทีอาร์เอ็นเอ – Thai" lang="th" hreflang="th" data-title="ทีอาร์เอ็นเอ" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Ta%C5%9F%C4%B1y%C4%B1c%C4%B1_RNA" title="Taşıyıcı RNA – Turkish" lang="tr" hreflang="tr" data-title="Taşıyıcı RNA" data-language-autonym="Türkçe" data-language-local-name="Turkish" 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src="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Peptide_syn.svg/220px-Peptide_syn.svg.png" decoding="async" width="220" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Peptide_syn.svg/330px-Peptide_syn.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/42/Peptide_syn.svg/440px-Peptide_syn.svg.png 2x" data-file-width="512" data-file-height="314" /></a><figcaption>The interaction of tRNA and mRNA in protein synthesis.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above summary">tRNA</th></tr><tr><th colspan="2" class="infobox-header" style="background-color:#dddddd;">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="white-space: nowrap;">Symbol</th><td class="infobox-data">t</td></tr><tr><th scope="row" class="infobox-label" style="white-space: nowrap;"><a href="/wiki/Rfam" title="Rfam">Rfam</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://rfam.org/family/RF00005">RF00005</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color:#dddddd;">Other data</th></tr><tr><th scope="row" class="infobox-label" style="white-space: nowrap;"><a href="/wiki/RNA" title="RNA">RNA</a> type</th><td class="infobox-data"><a href="/wiki/Gene" title="Gene">gene</a>, <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a></td></tr><tr><th scope="row" class="infobox-label" style="white-space: nowrap;"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> structures</th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://www.ebi.ac.uk/pdbe/entry/search/index/?searchParams=%7B%22rfam_accession%22:%5B%7B%22value%22:%22RF00005%22,%22condition1%22:%22AND%22,%22condition2%22:%22Equal%20to%22%7D%5D,%22resultState%22:%7B%22tabIndex%22:0,%22paginationIndex%22:1,%22perPage%22:%2210%22,%22sortBy%22:%22release_date%20desc%22%7D%7D">PDBe</a> 3icq, 1asy, 1asz, 1il2, 2tra, 3tra, 486d, 1fir, 1yfg, 3eph, 3epj, 3epk, 3epl, 1efw, 1c0a, 2ake, 2azx, 2dr2, 1f7u, 1f7v, 3foz, 2hgp, 2j00, 2j02, 2ow8, 2v46, 2v48, 2wdg, 2wdh, 2wdk, 2wdm, 2wh1</td></tr></tbody></table> <p><b>Transfer RNA</b> (abbreviated <b>tRNA</b> and formerly referred to as <b>sRNA</b>, for <b>soluble RNA</b><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>) is an adaptor <a href="/wiki/Molecule" title="Molecule">molecule</a> composed of <a href="/wiki/RNA" title="RNA">RNA</a>, typically 76 to 90 <a href="/wiki/Nucleotides" class="mw-redirect" title="Nucleotides">nucleotides</a> in length (in eukaryotes).<sup id="cite_ref-sharp1985_2-0" class="reference"><a href="#cite_note-sharp1985-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In a <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>, it provides the physical link between the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> in <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> (mRNA) and the <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> sequence of proteins, carrying the correct sequence of amino acids to be combined by the protein-synthesizing machinery, the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>. Each three-nucleotide <a href="/wiki/Codon" class="mw-redirect" title="Codon">codon</a> in mRNA is <a href="/wiki/Complementary_DNA" title="Complementary DNA">complemented</a> by a three-nucleotide <a href="#Anticodon">anticodon</a> in tRNA. As such, tRNAs are a necessary component of <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, the biological synthesis of new <a href="/wiki/Protein" title="Protein">proteins</a> in accordance with the genetic code. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=1" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The process of <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> starts with the information stored in the nucleotide sequence of <a href="/wiki/DNA" title="DNA">DNA</a>. This is first transformed into mRNA, then tRNA specifies which three-nucleotide codon from the genetic code corresponds to which amino acid.<sup id="cite_ref-crick_3-0" class="reference"><a href="#cite_note-crick-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Each mRNA codon is recognized by a particular type of tRNA, which docks to it along a three-nucleotide <a href="#Anticodon">anticodon</a>, and together they form three <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a> <a href="/wiki/Base_pair" title="Base pair">base pairs</a>. </p><p>On the other end of the tRNA is a covalent attachment to the amino acid corresponding to the anticodon sequence, with each type of tRNA attaching to a specific amino acid. Because the genetic code contains multiple codons that specify the same amino acid, there are several tRNA molecules bearing different anticodons which carry the same amino acid. </p><p>The covalent attachment to the tRNA <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">3' end</a> is catalysed by enzymes called <a href="/wiki/Aminoacyl_tRNA_synthetase" title="Aminoacyl tRNA synthetase">aminoacyl tRNA synthetases</a>. During protein synthesis, tRNAs with attached amino acids are delivered to the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> by proteins called <a href="/wiki/Elongation_factor" title="Elongation factor">elongation factors</a>, which aid in association of the tRNA with the ribosome, synthesis of the new polypeptide, and translocation (movement) of the ribosome along the mRNA. If the tRNA's anticodon matches the mRNA, another tRNA already <a href="#Binding_to_ribosome">bound to the ribosome</a> transfers the growing polypeptide chain from its 3' end to the amino acid attached to the 3' end of the newly delivered tRNA, a reaction catalysed by the ribosome. A large number of the individual nucleotides in a tRNA molecule may be <a href="/wiki/Chemical_modification#Chemical_modification_in_biochemistry" title="Chemical modification">chemically modified</a>, often by <a href="/wiki/Methylation" title="Methylation">methylation</a> or <a href="/wiki/Deamidation" title="Deamidation">deamidation</a>. These unusual bases sometimes affect the tRNA's interaction with <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a> and sometimes occur in the <a href="/wiki/Anticodon" class="mw-redirect" title="Anticodon">anticodon</a> to alter base-pairing properties.<sup id="cite_ref-Stryer2002_4-0" class="reference"><a href="#cite_note-Stryer2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=2" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:TRNA-Phe_yeast_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/TRNA-Phe_yeast_en.svg/220px-TRNA-Phe_yeast_en.svg.png" decoding="async" width="220" height="274" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/TRNA-Phe_yeast_en.svg/330px-TRNA-Phe_yeast_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/59/TRNA-Phe_yeast_en.svg/440px-TRNA-Phe_yeast_en.svg.png 2x" data-file-width="620" data-file-height="771" /></a><figcaption>Secondary cloverleaf structure of tRNA</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:TRNA-Phe_yeast_1ehz.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/TRNA-Phe_yeast_1ehz.png/220px-TRNA-Phe_yeast_1ehz.png" decoding="async" width="220" height="218" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/TRNA-Phe_yeast_1ehz.png/330px-TRNA-Phe_yeast_1ehz.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/ba/TRNA-Phe_yeast_1ehz.png/440px-TRNA-Phe_yeast_1ehz.png 2x" data-file-width="1174" data-file-height="1161" /></a><figcaption>Tertiary structure of tRNA. <span style="color:#E4D00A;"><i>CCA tail</i></span> in yellow, <span style="color:purple;"><i>acceptor stem</i></span> in purple, <span style="color:orange;"><i>variable loop</i></span> in orange, <span style="color:red;"><i>D arm</i></span> in red, <span style="color:blue;"><i>anticodon arm</i></span> in blue with <i>anticodon</i> in black, <span style="color:green;"><i>T arm</i></span> in green.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Trna.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Trna.gif/220px-Trna.gif" decoding="async" width="220" height="216" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Trna.gif/330px-Trna.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/b/b2/Trna.gif 2x" data-file-width="350" data-file-height="344" /></a><figcaption>3D animated GIF showing the structure of phenylalanine-tRNA from yeast (PDB ID 1ehz). White lines indicate base pairing by hydrogen bonds. In the orientation shown, the acceptor stem is on top and the anticodon on the bottom.<sup id="cite_ref-tRNA_proteopedia_5-0" class="reference"><a href="#cite_note-tRNA_proteopedia-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The structure of tRNA can be decomposed into its <a href="/wiki/Primary_structure" class="mw-redirect" title="Primary structure">primary structure</a>, its <a href="/wiki/Nucleic_acid_secondary_structure" title="Nucleic acid secondary structure">secondary structure</a> (usually visualized as the <i>cloverleaf structure</i>), and its <a href="/wiki/Tertiary_structure" class="mw-redirect" title="Tertiary structure">tertiary structure</a><sup id="cite_ref-itoh_6-0" class="reference"><a href="#cite_note-itoh-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> (all tRNAs have a similar L-shaped 3D structure that allows them to fit into the <a href="/wiki/P-site" title="P-site">P</a> and <a href="/wiki/A-site" title="A-site">A</a> sites of the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>). The cloverleaf structure becomes the 3D L-shaped structure through coaxial stacking of the helices, which is a common <a href="/wiki/Nucleic_acid_tertiary_structure" title="Nucleic acid tertiary structure">RNA tertiary structure</a> motif. The lengths of each arm, as well as the loop 'diameter', in a tRNA molecule vary from species to species.<sup id="cite_ref-itoh_6-1" class="reference"><a href="#cite_note-itoh-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-goodenbour2006_7-0" class="reference"><a href="#cite_note-goodenbour2006-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The tRNA structure consists of the following: </p> <ul><li>The <b>acceptor stem</b> is a 7- to 9-base pair (bp) stem made by the base pairing of the 5′-terminal nucleotide with the 3′-terminal nucleotide (which contains the CCA tail used to attach the amino acid). The acceptor stem may contain non-Watson-Crick base pairs.<sup id="cite_ref-itoh_6-2" class="reference"><a href="#cite_note-itoh-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li> <li>The <b>CCA tail</b> is a <a href="/wiki/Cytosine" title="Cytosine">cytosine</a>-cytosine-<a href="/wiki/Adenine" title="Adenine">adenine</a> sequence at the 3′ end of the tRNA molecule. The amino acid loaded onto the tRNA by <a href="/wiki/Aminoacyl_tRNA_synthetase" title="Aminoacyl tRNA synthetase">aminoacyl tRNA synthetases</a>, to form <a href="/wiki/Aminoacyl-tRNA" title="Aminoacyl-tRNA">aminoacyl-tRNA</a>, is covalently bonded to the 3′-hydroxyl group on the CCA tail.<sup id="cite_ref-ibba_9-0" class="reference"><a href="#cite_note-ibba-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> This sequence is important for the recognition of tRNA by enzymes and critical in translation.<sup id="cite_ref-pmid392600_10-0" class="reference"><a href="#cite_note-pmid392600-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9242921_11-0" class="reference"><a href="#cite_note-pmid9242921-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> In prokaryotes, the CCA sequence is transcribed in some tRNA sequences. In most prokaryotic tRNAs and eukaryotic tRNAs, the CCA sequence is added during processing and therefore does not appear in the tRNA gene.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup></li> <li>The <a href="/wiki/D_arm" title="D arm"><b>D loop</b></a> is a 4- to 6-bp stem ending in a loop that often contains <a href="/wiki/Dihydrouridine" title="Dihydrouridine">dihydrouridine</a>.<sup id="cite_ref-itoh_6-3" class="reference"><a href="#cite_note-itoh-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>The <b>anticodon loop</b> is a 5-bp stem whose loop contains the <a href="#Anticodon">anticodon</a>.<sup id="cite_ref-itoh_6-4" class="reference"><a href="#cite_note-itoh-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li> <li>The <a href="/wiki/T_arm" title="T arm"><b>TΨC loop</b></a> is named so because of the characteristic presence of the unusual base Ψ in the loop, where Ψ is <a href="/wiki/Pseudouridine" title="Pseudouridine">pseudouridine</a>, a modified <a href="/wiki/Uridine" title="Uridine">uridine</a>. The modified base is often found within the sequence 5'-TΨCGA-3', with the T (<a href="/wiki/Ribothymidine" class="mw-redirect" title="Ribothymidine">ribothymidine</a>, m5U) and A forming a base pair.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup></li> <li>The <b>variable loop</b> or <i>V loop</i> sits between the anticodon loop and the ΨU loop and, as its name implies, varies in size from 3 to 21 bases. In some tRNAs, the "loop" is long enough to form a rigid stem, the <i>variable arm</i>.<sup id="cite_ref-pmid35882385_14-0" class="reference"><a href="#cite_note-pmid35882385-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> tRNAs with a V loop more than 10 bases long is classified as "class II" and the rest is called "class I".<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Anticodon">Anticodon</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=3" title="Edit section: Anticodon"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An <b>anticodon</b><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> is a unit of three <a href="/wiki/Nucleotides" class="mw-redirect" title="Nucleotides">nucleotides</a> corresponding to the three bases of an <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> <a href="/wiki/Genetic_code" title="Genetic code">codon</a>. Each tRNA has a distinct anticodon triplet sequence that can form 3 <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a> <a href="/wiki/Base_pair" title="Base pair">base pairs</a> to one or more codons for an amino acid. Some anticodons pair with more than one codon due to <a href="/wiki/Wobble_base_pair" title="Wobble base pair">wobble base pairing</a>. Frequently, the first nucleotide of the anticodon is one not found on mRNA: <a href="/wiki/Inosine" title="Inosine">inosine</a>, which can <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bond</a> to more than one base in the corresponding codon position.<sup id="cite_ref-Stryer2002_4-1" class="reference"><a href="#cite_note-Stryer2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 29.3.9">: 29.3.9 </span></sup> In <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>, it is common for a single amino acid to be specified by all four third-position possibilities, or at least by both <a href="/wiki/Pyrimidine" title="Pyrimidine">pyrimidines</a> and <a href="/wiki/Purine" title="Purine">purines</a>; for example, the amino acid <a href="/wiki/Glycine" title="Glycine">glycine</a> is coded for by the codon sequences GGU, GGC, GGA, and GGG. Other modified nucleotides may also appear at the first anticodon position—sometimes known as the "wobble position"—resulting in subtle changes to the genetic code, as for example in <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> The possibility of wobble bases reduces the number of tRNA types required: instead of 61 types with one for each sense codon of the standard genetic code), only 31 tRNAs are required to translate, unambiguously, all 61 sense codons.<sup id="cite_ref-crick_3-1" class="reference"><a href="#cite_note-crick-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Nomenclature">Nomenclature</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=4" title="Edit section: Nomenclature"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A tRNA is commonly named by its intended amino acid (e.g. <span class="nowrap">tRNA-Asn</span>), by its anticodon sequence (e.g. <span class="nowrap">tRNA(GUU)</span>), or by both (e.g. <span class="nowrap">tRNA-Asn(GUU)</span> or <span class="nowrap">tRNA<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:-0.4em;line-height:1.2em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">Asn</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline">GUU</sub></span></span></span>).<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> These two features describe the main function of the tRNA, but do not actually cover the whole diversity of tRNA variation; as a result, numerical suffixes are added to differentiate.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> tRNAs intended for the same amino acid are called "isotypes"; these with the same anticodon sequence are called "isoacceptors"; and these with both being the same but differing in other places are called "isodecoders".<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Aminoacylation">Aminoacylation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=5" title="Edit section: Aminoacylation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Aminoacyl-tRNA" title="Aminoacyl-tRNA">Aminoacyl-tRNA</a></div> <p><a href="/wiki/Aminoacylation" title="Aminoacylation">Aminoacylation</a> is the process of adding an aminoacyl group to a compound. It covalently links an <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> to the CCA 3′ end of a tRNA molecule. Each tRNA is aminoacylated (or <i>charged</i>) with a specific amino acid by an <a href="/wiki/Aminoacyl_tRNA_synthetase" title="Aminoacyl tRNA synthetase">aminoacyl tRNA synthetase</a>. There is normally a single aminoacyl tRNA synthetase for each amino acid, despite the fact that there can be more than one tRNA, and more than one anticodon for an amino acid. Recognition of the appropriate tRNA by the synthetases is not mediated solely by the anticodon, and the acceptor stem often plays a prominent role.<sup id="cite_ref-pmid7692438_22-0" class="reference"><a href="#cite_note-pmid7692438-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Reaction: </p> <ol><li>amino acid + <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> → aminoacyl-AMP + <a href="/wiki/Pyrophosphate" title="Pyrophosphate">PPi</a></li> <li>aminoacyl-AMP + tRNA → aminoacyl-tRNA + <a href="/wiki/Adenosine_monophosphate" title="Adenosine monophosphate">AMP</a></li></ol> <p>Certain organisms can have one or more aminophosphate-tRNA synthetases missing. This leads to charging of the tRNA by a chemically related amino acid, and by use of an enzyme or enzymes, the tRNA is modified to be correctly charged. For example, <i><a href="/wiki/Helicobacter_pylori" title="Helicobacter pylori">Helicobacter pylori</a></i> has glutaminyl tRNA synthetase missing. Thus, glutamate tRNA synthetase charges tRNA-glutamine(tRNA-Gln) with <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>. An amidotransferase then converts the acid side chain of the glutamate to the amide, forming the correctly charged gln-tRNA-Gln. </p> <div class="mw-heading mw-heading2"><h2 id="Binding_to_ribosome">Binding to ribosome</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=6" title="Edit section: Binding to ribosome"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Transfer_RNA_morph_AT_to_PE_conformation.ogv/220px--Transfer_RNA_morph_AT_to_PE_conformation.ogv.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="147" data-durationhint="7" data-mwtitle="Transfer_RNA_morph_AT_to_PE_conformation.ogv" data-mwprovider="wikimediacommons" resource="/wiki/File:Transfer_RNA_morph_AT_to_PE_conformation.ogv"><source src="//upload.wikimedia.org/wikipedia/commons/7/7b/Transfer_RNA_morph_AT_to_PE_conformation.ogv" type="video/ogg; codecs="theora"" data-width="480" data-height="320" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/7/7b/Transfer_RNA_morph_AT_to_PE_conformation.ogv/Transfer_RNA_morph_AT_to_PE_conformation.ogv.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="480" data-height="320" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/7/7b/Transfer_RNA_morph_AT_to_PE_conformation.ogv/Transfer_RNA_morph_AT_to_PE_conformation.ogv.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="360" data-height="240" /></video></span><figcaption>The range of conformations adopted by tRNA as it transits the A/T through P/E sites on the ribosome. The Protein Data Bank (PDB) codes for the structural models used as end points of the animation are given. Both tRNAs are modeled as phenylalanine-specific tRNA from <i>Escherichia coli</i>, with the A/T tRNA as a homology model of the deposited coordinates. Color coding as shown for <a href="#Structure">tRNA tertiary structure</a>. Adapted from.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> has three binding sites for tRNA molecules that span the space between the two <a href="/wiki/Ribosome#Structure" title="Ribosome">ribosomal subunits</a>: the <a href="/wiki/A-site" title="A-site">A (aminoacyl)</a>,<sup id="cite_ref-pmid14681588_24-0" class="reference"><a href="#cite_note-pmid14681588-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> <a href="/wiki/P-site" title="P-site">P (peptidyl)</a>, and <a href="/wiki/E-site" title="E-site">E (exit) sites</a>. In addition, the ribosome has two other sites for tRNA binding that are used during <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> decoding or during the initiation of <a href="/wiki/Translation_(biology)" title="Translation (biology)">protein synthesis</a>. These are the T site (named <a href="/wiki/EF-Tu" title="EF-Tu">elongation factor Tu</a>) and I site (initiation).<sup id="cite_ref-Agirrezabala_25-0" class="reference"><a href="#cite_note-Agirrezabala-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Allen_26-0" class="reference"><a href="#cite_note-Allen-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> By convention, the tRNA binding sites are denoted with the site on the <a href="/wiki/Ribosome#Structure" title="Ribosome">small ribosomal subunit</a> listed first and the site on the <a href="/wiki/Ribosome#Structure" title="Ribosome">large ribosomal subunit</a> listed second. For example, the A site is often written A/A, the P site, P/P, and the E site, E/E.<sup id="cite_ref-Agirrezabala_25-1" class="reference"><a href="#cite_note-Agirrezabala-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> The binding proteins like L27, L2, L14, L15, L16 at the A- and P- sites have been determined by affinity labeling by A. P. Czernilofsky et al. (<i>Proc. Natl. Acad. Sci, USA</i>, pp. 230–234, 1974). </p><p>Once translation initiation is complete, the first aminoacyl tRNA is located in the P/P site, ready for the elongation cycle described below. During translation elongation, tRNA first binds to the ribosome as part of a complex with elongation factor Tu (<a href="/wiki/EF-Tu" title="EF-Tu">EF-Tu</a>) or its eukaryotic (<a href="/wiki/EEF-1" title="EEF-1">eEF-1</a>) or archaeal counterpart. This initial tRNA binding site is called the A/T site. In the A/T site, the A-site half resides in the <a href="/wiki/Ribosome#Structure" title="Ribosome">small ribosomal subunit</a> where the mRNA decoding site is located. The mRNA decoding site is where the <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> <a href="/wiki/Codon" class="mw-redirect" title="Codon">codon</a> is read out during translation. The T-site half resides mainly on the <a href="/wiki/Ribosome#Structure" title="Ribosome">large ribosomal subunit</a> where EF-Tu or eEF-1 interacts with the ribosome. Once mRNA decoding is complete, the aminoacyl-tRNA is bound in the A/A site and is ready for the next <a href="/wiki/Bacterial_translation#Elongation" title="Bacterial translation">peptide bond</a><sup id="cite_ref-PTC_27-0" class="reference"><a href="#cite_note-PTC-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> to be formed to its attached amino acid. The peptidyl-tRNA, which transfers the growing polypeptide to the aminoacyl-tRNA bound in the A/A site, is bound in the P/P site. Once the peptide bond is formed, the tRNA in the P/P site is acylated, or has a <a href="#Structure">free 3' end</a>, and the tRNA in the A/A site dissociates the growing polypeptide chain. To allow for the next elongation cycle, the tRNAs then move through hybrid A/P and P/E binding sites, before completing the cycle and residing in the P/P and E/E sites. Once the A/A and P/P tRNAs have moved to the P/P and E/E sites, the mRNA has also moved over by one <a href="/wiki/Codon" class="mw-redirect" title="Codon">codon</a> and the A/T site is vacant, ready for the next round of mRNA decoding. The tRNA bound in the E/E site then leaves the ribosome. </p><p>The P/I site is actually the first to bind to aminoacyl tRNA, which is delivered by an initiation factor called <a href="/wiki/Prokaryotic_initiation_factor-2" class="mw-redirect" title="Prokaryotic initiation factor-2">IF2</a> in bacteria.<sup id="cite_ref-Allen_26-1" class="reference"><a href="#cite_note-Allen-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> However, the existence of the P/I site in eukaryotic or archaeal <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a> has not yet been confirmed. The P-site protein L27 has been determined by affinity labeling by E. Collatz and A. P. Czernilofsky (<i>FEBS Lett.</i>, Vol. 63, pp. 283–286, 1976). </p> <div class="mw-heading mw-heading2"><h2 id="tRNA_genes">tRNA genes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=7" title="Edit section: tRNA genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Organisms vary in the number of tRNA <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> in their <a href="/wiki/Genome" title="Genome">genome</a>. For example, the <a href="/wiki/Nematode" title="Nematode">nematode</a> worm <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i>, a commonly used model organism in <a href="/wiki/Genetics" title="Genetics">genetics</a> studies, has 29,647 genes in its <a href="/wiki/Cell_nucleus" title="Cell nucleus">nuclear</a> genome,<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> of which 620 code for tRNA.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> The budding yeast <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> has 275 tRNA genes in its genome. The number of tRNA genes per genome can vary widely, with bacterial species from groups such as Fusobacteria and Tenericutes having around 30 genes per genome while complex eukaryotic genomes such as the zebrafish (<i>Danio rerio</i>) can bear more than 10 thousand tRNA genes.<sup id="cite_ref-:0_31-0" class="reference"><a href="#cite_note-:0-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>In the human genome, which, according to January 2013 estimates, has about 20,848 protein coding genes <sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> in total, there are 497 nuclear genes encoding cytoplasmic tRNA molecules, and 324 tRNA-derived <a href="/wiki/Pseudogenes" class="mw-redirect" title="Pseudogenes">pseudogenes</a>—tRNA genes thought to be no longer functional<sup id="cite_ref-Lander_33-0" class="reference"><a href="#cite_note-Lander-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> (although pseudo tRNAs have been shown to be involved in <a href="/wiki/Antimicrobial_resistance" title="Antimicrobial resistance">antibiotic resistance</a> in bacteria).<sup id="cite_ref-Rogers12_34-0" class="reference"><a href="#cite_note-Rogers12-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> As with all eukaryotes, there are 22 <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondrial</a> tRNA genes<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> in humans. Mutations in some of these genes have been associated with severe diseases like the <a href="/wiki/MELAS_syndrome" title="MELAS syndrome">MELAS syndrome</a>. Regions in nuclear <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a>, very similar in sequence to mitochondrial tRNA genes, have also been identified (tRNA-lookalikes).<sup id="cite_ref-Telonis14_36-0" class="reference"><a href="#cite_note-Telonis14-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> These tRNA-lookalikes are also considered part of the <a href="/wiki/Numt" class="mw-redirect" title="Numt">nuclear mitochondrial DNA</a> (genes transferred from the mitochondria to the nucleus).<sup id="cite_ref-Telonis14_36-1" class="reference"><a href="#cite_note-Telonis14-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Ramos11_37-0" class="reference"><a href="#cite_note-Ramos11-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The phenomenon of multiple nuclear copies of mitochondrial tRNA (tRNA-lookalikes) has been observed in many higher organisms from human to the opossum<sup id="cite_ref-Telonis15-look_38-0" class="reference"><a href="#cite_note-Telonis15-look-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> suggesting the possibility that the lookalikes are functional. </p><p>Cytoplasmic tRNA genes can be grouped into 49 families according to their anticodon features. These genes are found on all chromosomes, except the 22 and Y chromosome. High clustering on 6p is observed (140 tRNA genes), as well as on chromosome 1.<sup id="cite_ref-Lander_33-1" class="reference"><a href="#cite_note-Lander-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p><p>The <a href="/wiki/HUGO_Gene_Nomenclature_Committee" title="HUGO Gene Nomenclature Committee">HGNC</a>, in collaboration with the Genomic tRNA Database (<a rel="nofollow" class="external text" href="http://gtrnadb.ucsc.edu/">GtRNAdb</a>) and experts in the field, has approved unique names for human genes that encode tRNAs. </p><p>Typically, tRNAs genes from Bacteria are shorter (mean = 77.6 bp) than tRNAs from Archaea (mean = 83.1 bp) and eukaryotes (mean = 84.7 bp).<sup id="cite_ref-:0_31-1" class="reference"><a href="#cite_note-:0-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> The mature tRNA follows an opposite pattern with tRNAs from Bacteria being usually longer (median = 77.6 nt) than tRNAs from Archaea (median = 76.8 nt), with eukaryotes exhibiting the shortest mature tRNAs (median = 74.5 nt).<sup id="cite_ref-:0_31-2" class="reference"><a href="#cite_note-:0-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Evolution">Evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=8" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Genomic tRNA content is a differentiating feature of genomes among biological domains of life: Archaea present the simplest situation in terms of genomic tRNA content with a uniform number of gene copies, Bacteria have an intermediate situation and Eukarya present the most complex situation.<sup id="cite_ref-evamaria_39-0" class="reference"><a href="#cite_note-evamaria-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Eukarya present not only more tRNA gene content than the other two kingdoms but also a high variation in <a href="/wiki/Gene_copy_number" class="mw-redirect" title="Gene copy number">gene copy number</a> among different isoacceptors, and this complexity seem to be due to duplications of tRNA genes and changes in anticodon specificity <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (September 2015)">citation needed</span></a></i>]</sup>. </p><p>Evolution of the tRNA gene copy number across different species has been linked to the appearance of specific tRNA modification enzymes (uridine methyltransferases in Bacteria, and adenosine deaminases in Eukarya), which increase the decoding capacity of a given tRNA.<sup id="cite_ref-evamaria_39-1" class="reference"><a href="#cite_note-evamaria-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> As an example, tRNA<sup>Ala</sup> encodes four different tRNA isoacceptors (AGC, UGC, GGC and CGC). In Eukarya, AGC isoacceptors are extremely enriched in gene copy number in comparison to the rest of isoacceptors, and this has been correlated with its A-to-I modification of its wobble base. This same trend has been shown for most amino acids of eukaryal species. Indeed, the effect of these two tRNA modifications is also seen in <a href="/wiki/Codon_usage_bias" title="Codon usage bias">codon usage bias</a>. Highly expressed genes seem to be enriched in codons that are exclusively using codons that will be decoded by these modified tRNAs, which suggests a possible role of these codons—and consequently of these tRNA modifications—in translation efficiency.<sup id="cite_ref-evamaria_39-2" class="reference"><a href="#cite_note-evamaria-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>Many species have lost specific tRNAs during evolution. For instance, both mammals and birds lack the same 14 out of the possible 64 tRNA genes, but other life forms contain these tRNAs.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> For translating codons for which an exactly pairing tRNA is missing, organisms resort to a strategy called <a href="/wiki/Wobble_base_pair" title="Wobble base pair">wobbling</a>, in which imperfectly matched tRNA/mRNA pairs still give rise to translation, although this strategy also increases the propensity for translation errors.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> The reasons why tRNA genes have been lost during evolution remains under debate but may relate improving resistance to viral infection.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Because nucleotide triplets can present more combinations than there are amino acids and associated tRNAs, there is redundancy in the genetic code, and several different 3-nucleotide codons can express the same amino acid. This codon bias is what necessitates codon optimization. </p> <div class="mw-heading mw-heading4"><h4 id="Hypothetical_origin">Hypothetical origin</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=9" title="Edit section: Hypothetical origin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The top half of tRNA (consisting of the T arm and the acceptor stem with 5′-terminal phosphate group and 3′-terminal CCA group) and the bottom half (consisting of the D arm and the anticodon arm) are independent units in structure as well as in function. The top half may have evolved first including the 3′-terminal genomic tag which originally may have marked tRNA-like molecules for replication in early <a href="/wiki/RNA_world" title="RNA world">RNA world</a>. The bottom half may have evolved later as an expansion, e.g. as protein synthesis started in RNA world and turned it into a ribonucleoprotein world (<a href="/wiki/RNP_world" title="RNP world">RNP world</a>). This proposed scenario is called <a href="/w/index.php?title=Genomic_tag_hypothesis&action=edit&redlink=1" class="new" title="Genomic tag hypothesis (page does not exist)">genomic tag hypothesis</a>. In fact, tRNA and tRNA-like aggregates have an important catalytic influence (i.e., as <a href="/wiki/Ribozyme" title="Ribozyme">ribozymes</a>) on replication still today. These roles may be regarded as '<a href="/wiki/Molecular_fossil" class="mw-redirect" title="Molecular fossil">molecular (or chemical) fossils</a>' of RNA world.<sup id="cite_ref-MW_GenomicTag_43-0" class="reference"><a href="#cite_note-MW_GenomicTag-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> In March 2021, researchers reported evidence suggesting that an early form of transfer RNA could have been a replicator <a href="/wiki/Ribozyme" title="Ribozyme">ribozyme</a> molecule in the very early development of life, or <a href="/wiki/Abiogenesis" title="Abiogenesis">abiogenesis</a>.<sup id="cite_ref-EL-20210302_44-0" class="reference"><a href="#cite_note-EL-20210302-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-STD-20210403_45-0" class="reference"><a href="#cite_note-STD-20210403-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>Evolution of type I and type II tRNAs is explained to the last nucleotide by the three 31 nucleotide minihelix tRNA evolution theorem, which also describes the pre-life to life transition on Earth.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Three 31 nucleotide minihelices of known sequence were ligated in pre-life to generate a 93 nucleotide tRNA precursor. In pre-life, a 31 nucleotide D loop minihelix (GCGGCGGUAGCCUAGCCUAGCCUACCGCCGC) was ligated to two 31 nucleotide anticodon loop minihelices (GCGGCGGCCGGGCU/???AACCCGGCCGCCGC; / indicates a U-turn conformation in the RNA backbone; ? indicates unknown base identity) to form the 93 nucleotide tRNA precursor. To generate type II tRNAs, a single internal 9 nucleotide deletion occurred within ligated acceptor stems (CCGCCGCGCGGCGG goes to GGCGG). To generate type I tRNAs, an additional, related 9 nucleotide deletion occurred within ligated acceptor stems within the variable loop region (CCGCCGCGCGGCGG goes to CCGCC). These two 9 nucleotide deletions are identical on complementary RNA strands. tRNAomes (all of the tRNAs of an organism) were generated by duplication and mutation. </p><p>Very clearly, life evolved from a polymer world that included RNA repeats and RNA inverted repeats (stem-loop-stems). Of particular importance were the 7 nucleotide U-turn loops (CU/???AA). After LUCA (the last universal common (cellular) ancestor), the T loop evolved to interact with the D loop at the tRNA “elbow” (T loop: UU/CAAAU, after LUCA). Polymer world progressed to minihelix world to tRNA world, which has endured for ~4 billion years. Analysis of tRNA sequences reveals a major successful pathway in evolution of life on Earth. </p> <div class="mw-heading mw-heading3"><h3 id="tRNA-derived_fragments">tRNA-derived fragments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=10" title="Edit section: tRNA-derived fragments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>tRNA-derived fragments (or tRFs) are short molecules that emerge after cleavage of the mature tRNAs or the precursor transcript.<sup id="cite_ref-Gebetsberger13_51-0" class="reference"><a href="#cite_note-Gebetsberger13-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Shigematsu14_52-0" class="reference"><a href="#cite_note-Shigematsu14-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Sobala11_53-0" class="reference"><a href="#cite_note-Sobala11-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Keam15_54-0" class="reference"><a href="#cite_note-Keam15-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Both cytoplasmic and mitochondrial tRNAs can produce fragments.<sup id="cite_ref-Telonis15-dissect_55-0" class="reference"><a href="#cite_note-Telonis15-dissect-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> There are at least four structural types of tRFs believed to originate from mature tRNAs, including the relatively long tRNA halves and short 5'-tRFs, 3'-tRFs and i-tRFs.<sup id="cite_ref-Gebetsberger13_51-1" class="reference"><a href="#cite_note-Gebetsberger13-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Telonis15-dissect_55-1" class="reference"><a href="#cite_note-Telonis15-dissect-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kumar14_56-0" class="reference"><a href="#cite_note-Kumar14-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> The precursor tRNA can be cleaved to produce molecules from the 5' leader or 3' trail sequences. Cleavage enzymes include Angiogenin, Dicer, RNase Z and RNase P.<sup id="cite_ref-Gebetsberger13_51-2" class="reference"><a href="#cite_note-Gebetsberger13-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Shigematsu14_52-1" class="reference"><a href="#cite_note-Shigematsu14-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Especially in the case of Angiogenin, the tRFs have a characteristically unusual cyclic phosphate at their 3' end and a hydroxyl group at the 5' end.<sup id="cite_ref-Honda15_57-0" class="reference"><a href="#cite_note-Honda15-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> tRFs appear to play a role in <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a>, specifically in the suppression of retroviruses and retrotransposons that use tRNA as a primer for replication. Half-tRNAs cleaved by <a href="/wiki/Angiogenin" title="Angiogenin">angiogenin</a> are also known as tiRNAs. The biogenesis of smaller fragments, including those that function as <a href="/wiki/PiRNA" class="mw-redirect" title="PiRNA">piRNAs</a>, are less understood.<sup id="cite_ref-pmid29934075_58-0" class="reference"><a href="#cite_note-pmid29934075-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p><p>tRFs have multiple dependencies and roles; such as exhibiting significant changes between sexes, among races and disease status.<sup id="cite_ref-Telonis15-dissect_55-2" class="reference"><a href="#cite_note-Telonis15-dissect-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Telonis18_59-0" class="reference"><a href="#cite_note-Telonis18-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Telonis19_60-0" class="reference"><a href="#cite_note-Telonis19-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Functionally, they can be loaded on Ago and act through RNAi pathways,<sup id="cite_ref-Sobala11_53-1" class="reference"><a href="#cite_note-Sobala11-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kumar14_56-1" class="reference"><a href="#cite_note-Kumar14-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Shigematsu15_61-0" class="reference"><a href="#cite_note-Shigematsu15-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> participate in the formation of stress granules,<sup id="cite_ref-Emara10_62-0" class="reference"><a href="#cite_note-Emara10-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> displace mRNAs from RNA-binding proteins<sup id="cite_ref-Goodarzi15_63-0" class="reference"><a href="#cite_note-Goodarzi15-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> or inhibit translation.<sup id="cite_ref-Ivanov11_64-0" class="reference"><a href="#cite_note-Ivanov11-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> At the system or the organismal level, the four types of tRFs have a diverse spectrum of activities. Functionally, tRFs are associated with viral infection,<sup id="cite_ref-Selitsky15_65-0" class="reference"><a href="#cite_note-Selitsky15-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> cancer,<sup id="cite_ref-Kumar14_56-2" class="reference"><a href="#cite_note-Kumar14-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> cell proliferation <sup id="cite_ref-Honda15_57-1" class="reference"><a href="#cite_note-Honda15-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> and also with epigenetic transgenerational regulation of metabolism.<sup id="cite_ref-Sharma16_66-0" class="reference"><a href="#cite_note-Sharma16-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p><p>tRFs are not restricted to humans and have been shown to exist in multiple organisms.<sup id="cite_ref-Kumar14_56-3" class="reference"><a href="#cite_note-Kumar14-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Casas15_67-0" class="reference"><a href="#cite_note-Casas15-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hirose15_68-0" class="reference"><a href="#cite_note-Hirose15-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Karaiskos15_69-0" class="reference"><a href="#cite_note-Karaiskos15-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p><p>Two online tools are available for those wishing to learn more about tRFs: the framework for the interactive exploration of <u>mi</u>tochondrial and <u>n</u>uclear <u>t</u>RNA fragments (<a rel="nofollow" class="external text" href="https://cm.jefferson.edu/MINTbase/">MINTbase</a>)<sup id="cite_ref-Pliatsika16_70-0" class="reference"><a href="#cite_note-Pliatsika16-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pliatsika18_71-0" class="reference"><a href="#cite_note-Pliatsika18-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> and the relational database of <u>T</u>ransfer <u>R</u>NA related <u>F</u>ragments (<a rel="nofollow" class="external text" href="http://genome.bioch.virginia.edu/trfdb/">tRFdb</a>).<sup id="cite_ref-Kumar15_72-0" class="reference"><a href="#cite_note-Kumar15-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> MINTbase also provides a naming scheme for the naming of tRFs called <a rel="nofollow" class="external text" href="https://cm.jefferson.edu/MINTcodes/">tRF-license plates</a> (or MINTcodes) that is genome independent; the scheme compresses an RNA sequence into a shorter string. </p> <div class="mw-heading mw-heading3"><h3 id="Engineered_tRNAs">Engineered tRNAs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=11" title="Edit section: Engineered tRNAs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Genetic_code_expansion" class="mw-redirect" title="Genetic code expansion">Genetic code expansion</a></div> <p>tRNAs with modified anticodons and/or acceptor stems can be used to modify the genetic code. Scientists have successfully repurposed codons (sense and stop) to accept amino acids (natural and novel), for both initiation (see: <a href="/wiki/Start_codon" title="Start codon">start codon</a>) and elongation. </p><p>In 1990, tRNA<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:-0.4em;line-height:1.2em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">fMet2</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline">CUA</sub></span></span> (modified from the tRNA<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:-0.4em;line-height:1.2em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">fMet2</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline">CAU</sub></span></span> gene <a rel="nofollow" class="external text" href="https://ecocyc.org/gene?orgid=ECOLI&id=EG30061">metY</a>) was inserted into <i>E. coli</i>, causing it to initiate protein synthesis at the UAG stop codon, as long as it is preceded by a strong <a href="/wiki/Shine-Dalgarno_sequence" class="mw-redirect" title="Shine-Dalgarno sequence">Shine-Dalgarno sequence</a>. At initiation it not only inserts the traditional <a href="/wiki/Formylmethionine" class="mw-redirect" title="Formylmethionine">formylmethionine</a>, but also formylglutamine, as glutamyl-tRNA synthase also recognizes the new tRNA.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> The experiment was repeated in 1993, now with an elongator tRNA modified to be recognized by the <a href="/wiki/Methionyl-tRNA_formyltransferase" title="Methionyl-tRNA formyltransferase">methionyl-tRNA formyltransferase</a>.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> A similar result was obtained in <i><a href="/wiki/Mycobacterium" title="Mycobacterium">Mycobacterium</a></i>.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> Later experiments showed that the new tRNA was orthogonal to the regular AUG start codon showing no detectable off-target translation initiation events in a genomically recoded <i>E. coli</i> strain.<sup id="cite_ref-fmet2-cua-ec_76-0" class="reference"><a href="#cite_note-fmet2-cua-ec-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="tRNA_biogenesis">tRNA biogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=12" title="Edit section: tRNA biogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> cells, tRNAs are <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a> by <a href="/wiki/RNA_polymerase_III" title="RNA polymerase III">RNA polymerase III</a> as pre-tRNAs in the nucleus.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> RNA polymerase III recognizes two highly conserved downstream promoter sequences: the 5′ intragenic control region (5′-ICR, D-control region, or A box), and the 3′-ICR (T-control region or B box) inside tRNA genes.<sup id="cite_ref-sharp1985_2-1" class="reference"><a href="#cite_note-sharp1985-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-sharp1982_78-0" class="reference"><a href="#cite_note-sharp1982-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dieci_79-0" class="reference"><a href="#cite_note-Dieci-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> The first promoter begins at +8 of mature tRNAs and the second promoter is located 30–60 nucleotides downstream of the first promoter. The transcription terminates after a stretch of four or more <a href="/wiki/Thymidine" title="Thymidine">thymidines</a>.<sup id="cite_ref-sharp1985_2-2" class="reference"><a href="#cite_note-sharp1985-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dieci_79-1" class="reference"><a href="#cite_note-Dieci-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Bulge-helix-bulge_BHB_tRNA_intron.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Bulge-helix-bulge_BHB_tRNA_intron.png/220px-Bulge-helix-bulge_BHB_tRNA_intron.png" decoding="async" width="220" height="336" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Bulge-helix-bulge_BHB_tRNA_intron.png/330px-Bulge-helix-bulge_BHB_tRNA_intron.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Bulge-helix-bulge_BHB_tRNA_intron.png/440px-Bulge-helix-bulge_BHB_tRNA_intron.png 2x" data-file-width="552" data-file-height="844" /></a><figcaption>Bulge-helix-bulge motif of a tRNA intron</figcaption></figure> <p>Pre-tRNAs undergo extensive modifications inside the nucleus. Some pre-tRNAs contain <a href="/wiki/Intron" title="Intron">introns</a> that are spliced, or cut, to form the functional tRNA molecule;<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> in bacteria these self-<a href="/wiki/Splicing_(genetics)" class="mw-redirect" title="Splicing (genetics)">splice</a>, whereas in eukaryotes and <a href="/wiki/Archaea" title="Archaea">archaea</a> they are removed by tRNA-splicing <a href="/wiki/Endonuclease" title="Endonuclease">endonucleases</a>.<sup id="cite_ref-81" class="reference"><a href="#cite_note-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> Eukaryotic pre-tRNA contains bulge-helix-bulge (BHB) structure motif that is important for recognition and precise splicing of tRNA intron by endonucleases.<sup id="cite_ref-Soma2014_82-0" class="reference"><a href="#cite_note-Soma2014-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> This motif position and structure are evolutionarily conserved. However, some organisms, such as unicellular algae have a non-canonical position of BHB-motif as well as 5′- and 3′-ends of the spliced intron sequence.<sup id="cite_ref-Soma2014_82-1" class="reference"><a href="#cite_note-Soma2014-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> The 5′ sequence is removed by <a href="/wiki/RNase_P" class="mw-redirect" title="RNase P">RNase P</a>,<sup id="cite_ref-pmid9759486_83-0" class="reference"><a href="#cite_note-pmid9759486-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> whereas the 3′ end is removed by the <a href="/wiki/Ribonuclease_Z" title="Ribonuclease Z">tRNase Z</a> enzyme.<sup id="cite_ref-pmid17305600_84-0" class="reference"><a href="#cite_note-pmid17305600-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> A notable exception is in the <a href="/wiki/Archaeon" class="mw-redirect" title="Archaeon">archaeon</a> <i><a href="/wiki/Nanoarchaeum_equitans" title="Nanoarchaeum equitans">Nanoarchaeum equitans</a>,</i> which does not possess an RNase P enzyme and has a promoter placed such that transcription starts at the 5′ end of the mature tRNA.<sup id="cite_ref-pmid18451863_85-0" class="reference"><a href="#cite_note-pmid18451863-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> The non-templated 3′ CCA tail is added by a <a href="/wiki/Nucleotidyl_transferase" class="mw-redirect" title="Nucleotidyl transferase">nucleotidyl transferase</a>.<sup id="cite_ref-pmid15498478_86-0" class="reference"><a href="#cite_note-pmid15498478-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> Before tRNAs are <a href="/wiki/Nuclear_export" class="mw-redirect" title="Nuclear export">exported</a> into the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> by Los1/<a href="/wiki/Xpo-t" class="mw-redirect" title="Xpo-t">Xpo-t</a>,<sup id="cite_ref-pmid9660920_87-0" class="reference"><a href="#cite_note-pmid9660920-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid9512417_88-0" class="reference"><a href="#cite_note-pmid9512417-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> tRNAs are <a href="/wiki/Aminoacylation" title="Aminoacylation">aminoacylated</a>.<sup id="cite_ref-pmid9857198_89-0" class="reference"><a href="#cite_note-pmid9857198-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> The order of the processing events is not conserved. For example, in <a href="/wiki/Yeast" title="Yeast">yeast</a>, the splicing is not carried out in the nucleus but at the cytoplasmic side of <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondrial</a> membranes.<sup id="cite_ref-pmid12925762_90-0" class="reference"><a href="#cite_note-pmid12925762-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=13" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The existence of tRNA was first hypothesized by <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> as the "<a href="/wiki/Adaptor_hypothesis" title="Adaptor hypothesis">adaptor hypothesis</a>" based on the assumption that there must exist an adapter molecule capable of mediating the translation of the RNA alphabet into the protein alphabet. <a href="/wiki/Paul_C_Zamecnik" class="mw-redirect" title="Paul C Zamecnik">Paul C Zamecnik</a>, <a href="/wiki/Mahlon_Hoagland" title="Mahlon Hoagland">Mahlon Hoagland</a>, and <a href="/w/index.php?title=Mary_Louise_Stephenson&action=edit&redlink=1" class="new" title="Mary Louise Stephenson (page does not exist)">Mary Louise Stephenson</a> discovered tRNA. <sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> Significant research on structure was conducted in the early 1960s by <a href="/wiki/Alex_Rich" class="mw-redirect" title="Alex Rich">Alex Rich</a> and <a href="/wiki/Donald_Caspar" title="Donald Caspar">Donald Caspar</a>, two researchers in Boston, the Jacques Fresco group in <a href="/wiki/Princeton_University" title="Princeton University">Princeton University</a> and a <a href="/wiki/United_Kingdom" title="United Kingdom">United Kingdom</a> group at <a href="/wiki/King%27s_College_London" title="King's College London">King's College London</a>.<sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> In 1965, <a href="/wiki/Robert_W._Holley" title="Robert W. Holley">Robert W. Holley</a> of <a href="/wiki/Cornell_University" title="Cornell University">Cornell University</a> reported the primary structure and suggested three secondary structures.<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> tRNA was first crystallized in Madison, Wisconsin, by Robert M. Bock.<sup id="cite_ref-NYT1991_96-0" class="reference"><a href="#cite_note-NYT1991-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> The cloverleaf structure was ascertained by several other studies in the following years<sup id="cite_ref-97" class="reference"><a href="#cite_note-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> and was finally confirmed using <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a> studies in 1974. Two independent groups, <a href="/wiki/Kim_Sung-Hou" title="Kim Sung-Hou">Kim Sung-Hou</a> working under <a href="/wiki/Alexander_Rich" title="Alexander Rich">Alexander Rich</a> and a British group headed by <a href="/wiki/Aaron_Klug" title="Aaron Klug">Aaron Klug</a>, published the same crystallography findings within a year.<sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_relevance">Clinical relevance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=14" title="Edit section: Clinical relevance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Interference with aminoacylation may be useful as an approach to treating some diseases: cancerous cells may be relatively vulnerable to disturbed aminoacylation compared to healthy cells. The protein synthesis associated with cancer and viral biology is often very dependent on specific tRNA molecules. For instance, for liver cancer charging tRNA-Lys-CUU with lysine sustains liver cancer cell growth and metastasis, whereas healthy cells have a much lower dependence on this tRNA to support cellular physiology.<sup id="cite_ref-pmid33084231_100-0" class="reference"><a href="#cite_note-pmid33084231-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> Similarly, hepatitis E virus requires a tRNA landscape that substantially differs from that associated with uninfected cells.<sup id="cite_ref-pmid_32133647_101-0" class="reference"><a href="#cite_note-pmid_32133647-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> Hence, inhibition of aminoacylation of specific tRNA species is considered a promising novel avenue for the rational treatment of a plethora of diseases. </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Transfer_RNA&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 22em;"> <ul><li><a href="/wiki/Cloverleaf_model_of_tRNA" title="Cloverleaf model of tRNA">Cloverleaf model of tRNA</a></li> <li><a href="/wiki/Kim_Sung-Hou" title="Kim Sung-Hou">Kim Sung-Hou</a></li> <li><a href="/wiki/Kissing_stem-loop" title="Kissing stem-loop">Kissing stem-loop</a></li> <li><a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a></li> <li><a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding RNA</a> and <a href="/wiki/Intron" title="Intron">introns</a></li> <li><a href="/wiki/Slippery_sequence" title="Slippery sequence">Slippery sequence</a></li> <li><a href="/wiki/TmRNA" class="mw-redirect" title="TmRNA">tmRNA</a></li> <li><a href="/wiki/Transfer_RNA-like_structures" title="Transfer RNA-like structures">Transfer RNA-like structures</a></li> <li><a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">Translation</a></li> <li><a href="/wiki/TRNADB" title="TRNADB">tRNADB</a></li> <li><a href="/wiki/Wobble_hypothesis" class="mw-redirect" title="Wobble hypothesis">Wobble hypothesis</a></li> <li><a href="/wiki/Aminoacyl-tRNA" title="Aminoacyl-tRNA">Aminoacyl-tRNA</a></li></ul> </div> <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=Transfer_RNA&action=edit&section=16" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-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="CITEREFPlesciaPalczukCora-FigueroaMukherjee1965" class="citation journal cs1">Plescia OJ, Palczuk NC, Cora-Figueroa E, Mukherjee A, Braun W (October 1965). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC219862">"Production of antibodies to soluble RNA (sRNA)"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>54</b> (4): 1281–1285. <a 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</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=Transfer_RNA&action=edit&section=17" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output 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li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:TRNA" class="extiw" title="commons:Category:TRNA">TRNA</a></span>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://trnadb.bioinf.uni-leipzig.de/">tRNAdb (updated and completely restructured version of Spritzls tRNA compilation)</a></li> <li><a rel="nofollow" class="external text" href="https://www.the-scientist.com/news-opinion/transfer-rnas-have-a-surprising-role-in-breast-cancer-growth-70873">tRNA surprising role in breast cancer growth</a></li> <li><a rel="nofollow" class="external text" href="http://news.bbc.co.uk/2/hi/health/3762664.stm">tRNA link to heart disease and stroke</a></li> <li><a rel="nofollow" class="external text" href="http://gtrnadb.ucsc.edu/">GtRNAdb: Collection of tRNAs identified from complete genomes</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20141221113121/http://www.genenames.org/rna/TRNA">HGNC: Gene nomenclature of human tRNAs</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20091027143923/http://www.pdb.org/pdb/static.do?p=education_discussion%2Fmolecule_of_the_month%2Findex.html">Molecule of the Month</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070205000027/http://home.rcsb.org/">© RCSB Protein Data Bank</a>: <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100513191811/http://www.pdb.org/pdb/static.do?p=education_discussion%2Fmolecule_of_the_month%2Fpdb15_1.html">Transfer RNA</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100116025139/http://www.pdb.org/pdb/static.do?p=education_discussion%2Fmolecule_of_the_month%2Fpdb16_1.html">Aminoacyl-tRNA Synthetases</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20110316083658/http://www.pdb.org/pdb/static.do?p=education_discussion%2Fmolecule_of_the_month%2Fpdb81_1.html">Elongation Factors</a></li></ul></li> <li><a rel="nofollow" class="external text" href="http://rfam.org/family/RF00005">Rfam entry for tRNA</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist 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href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Nucleic_acids" title="Template:Nucleic acids"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nucleic_acids" title="Template talk:Nucleic acids"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nucleic_acids" title="Special:EditPage/Template:Nucleic acids"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Types_of_nucleic_acids" style="font-size:114%;margin:0 4em">Types of <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids </a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">Constituents</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/Nucleobase" class="mw-redirect" title="Nucleobase">Nucleobases</a></li> <li><a href="/wiki/Nucleoside" title="Nucleoside">Nucleosides</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotides</a></li> <li><a href="/wiki/Deoxyribonucleotide" title="Deoxyribonucleotide">Deoxynucleotides</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/RNA" title="RNA">Ribonucleic acids </a><br /><span class="nobold">(coding, <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding</a>)</span></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%;text-align: center;"><a href="/wiki/RNA#In_translation" title="RNA">Translational</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/Messenger_RNA" title="Messenger RNA">Messenger </a> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">precursor, heterogenous nuclear</a></li></ul></li> <li><a href="/wiki/Nucleoside-modified_messenger_RNA" title="Nucleoside-modified messenger RNA">modified Messenger</a></li> <li><a class="mw-selflink selflink">Transfer </a></li> <li><a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">Ribosomal </a></li> <li><a href="/wiki/Transfer-messenger_RNA" title="Transfer-messenger RNA">Transfer-messenger </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/RNA#Regulatory_RNAs" title="RNA">Regulatory</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/RNA_interference" title="RNA interference">Interferential </a> <ul><li><a href="/wiki/MicroRNA" title="MicroRNA">Micro </a></li> <li><a href="/wiki/Small_interfering_RNA" title="Small interfering RNA">Small interfering </a></li> <li><a href="/wiki/Piwi-interacting_RNA" title="Piwi-interacting RNA">Piwi-interacting</a></li></ul></li> <li><a href="/wiki/Antisense_RNA" title="Antisense RNA">Antisense </a></li> <li><a href="/wiki/RNA#In_RNA_processing" title="RNA">Processual</a> <ul><li><a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">Small nuclear </a></li> <li><a href="/wiki/Small_nucleolar_RNA" title="Small nucleolar RNA">Small nucleolar </a></li> <li><a href="/wiki/Small_Cajal_body-specific_RNA" title="Small Cajal body-specific RNA">Small Cajal Body RNAs</a></li> <li><a href="/wiki/Y_RNA" title="Y RNA">Y RNA</a></li></ul></li> <li><a href="/wiki/Enhancer_RNA" title="Enhancer RNA">Enhancer RNAs </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Others</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/Guide_RNA" title="Guide RNA">Guide </a></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Small_hairpin_RNA" class="mw-redirect" title="Small hairpin RNA">Small hairpin </a></li> <li><a href="/wiki/Small_temporal_RNA" title="Small temporal RNA">Small temporal </a></li> <li><a href="/wiki/Trans-acting_siRNA" title="Trans-acting siRNA">Trans-acting small interfering</a></li> <li><a href="/wiki/Subgenomic_mRNA" title="Subgenomic mRNA">Subgenomic messenger</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/DNA" title="DNA">Deoxyribonucleic <br />acids </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/Organellar_DNA" title="Organellar DNA">Organellar</a> <ul><li><a href="/wiki/Chloroplast_DNA" title="Chloroplast DNA">Chloroplast </a></li> <li><a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial</a></li></ul></li> <li><a href="/wiki/Complementary_DNA" title="Complementary DNA">Complementary </a></li> <li><a href="/wiki/Deoxyribozyme" title="Deoxyribozyme">Deoxyribozyme</a></li> <li><a href="/wiki/Genomic_DNA" title="Genomic DNA">Genomic </a></li> <li><a href="/wiki/Hachimoji_DNA" title="Hachimoji DNA">Hachimoji</a></li> <li><a href="/wiki/Multicopy_single-stranded_DNA" title="Multicopy single-stranded DNA">Multicopy single-stranded </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid_analogue" title="Nucleic acid analogue">Analogues</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/Xeno_nucleic_acid" title="Xeno nucleic acid">Xeno </a> <ul><li><a href="/wiki/Glycol_nucleic_acid" title="Glycol nucleic acid">Glycol </a></li> <li><a href="/wiki/Threose_nucleic_acid" title="Threose nucleic acid">Threose </a></li> <li><a href="/wiki/Hexose" title="Hexose">Hexose</a></li></ul></li> <li><a href="/wiki/Locked_nucleic_acid" title="Locked nucleic acid">Locked </a></li> <li><a href="/wiki/Peptide_nucleic_acid" title="Peptide nucleic acid">Peptide </a></li> <li><a href="/wiki/Morpholino" title="Morpholino">Morpholino</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Cloning_vector" title="Cloning vector">Cloning vectors</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/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Lambda_phage" title="Lambda phage">Lambda phage</a></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Artificial chromosomes</a> <ul><li><a href="/wiki/P1-derived_artificial_chromosome" title="P1-derived artificial chromosome">P1-derived </a></li> <li><a href="/wiki/Bacterial_artificial_chromosome" title="Bacterial artificial chromosome">Bacterial </a></li> <li><a href="/wiki/Yeast_artificial_chromosome" title="Yeast artificial chromosome">Yeast </a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Human</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align: center;"><div> <ul><li><span class="noviewer" 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title="Mitochondrion">Mitochondrial</a> proteins</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Outer_mitochondrial_membrane" class="mw-redirect" title="Outer mitochondrial membrane">Outer membrane</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%"><span class="nobold"><i><a href="/wiki/Fatty_acid_degradation" title="Fatty acid degradation">fatty acid degradation</a></i></span></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/Carnitine_palmitoyltransferase_I" title="Carnitine palmitoyltransferase I">Carnitine palmitoyltransferase I</a></li> <li><a href="/wiki/Long-chain-fatty-acid%E2%80%94CoA_ligase" title="Long-chain-fatty-acid—CoA ligase">Long-chain-fatty-acid—CoA ligase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> metabolism</i></span></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/Kynureninase" title="Kynureninase">Kynureninase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Monoamine_neurotransmitter" title="Monoamine neurotransmitter">monoamine neurotransmitter</a><br /> metabolism</i></span></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/Monoamine_oxidase" title="Monoamine oxidase">Monoamine oxidase</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/Mitochondrial_intermembrane_space" class="mw-redirect" title="Mitochondrial intermembrane space">Intermembrane space</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/Adenylate_kinase" title="Adenylate kinase">Adenylate kinase</a></li> <li><a href="/wiki/Creatine_kinase" title="Creatine kinase">Creatine kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">Inner membrane</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%"><span class="nobold"><i><a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a></i></span></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/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Coenzyme Q – cytochrome c reductase</a></li> <li><a href="/wiki/Cytochrome_c" title="Cytochrome c">Cytochrome c</a></li> <li><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a></li> <li><a href="/wiki/Succinate_dehydrogenase" title="Succinate dehydrogenase">Succinate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Pyrimidine_metabolism" title="Pyrimidine metabolism">pyrimidine metabolism</a></i></span></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/Dihydroorotate_dehydrogenase" title="Dihydroorotate dehydrogenase">Dihydroorotate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Mitochondrial_shuttle" title="Mitochondrial shuttle">mitochondrial shuttle</a></i></span></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/Malate-aspartate_shuttle" class="mw-redirect" title="Malate-aspartate shuttle">Malate-aspartate shuttle</a></li> <li><a href="/wiki/Glycerol_phosphate_shuttle" title="Glycerol phosphate shuttle">Glycerol phosphate shuttle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Steroidogenesis" class="mw-redirect" title="Steroidogenesis">steroidogenesis</a></i></span></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/Cholesterol_side-chain_cleavage_enzyme" title="Cholesterol side-chain cleavage enzyme">Cholesterol side-chain cleavage enzyme</a></li> <li><a href="/wiki/Steroid_11-beta-hydroxylase" class="mw-redirect" title="Steroid 11-beta-hydroxylase">Steroid 11-beta-hydroxylase</a></li> <li><a href="/wiki/Aldosterone_synthase" title="Aldosterone synthase">Aldosterone synthase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>other</i></span></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/Glutamate_aspartate_transporter" class="mw-redirect" title="Glutamate aspartate transporter">Glutamate aspartate transporter</a></li> <li><a href="/wiki/Glycerol-3-phosphate_dehydrogenase" title="Glycerol-3-phosphate dehydrogenase">Glycerol-3-phosphate dehydrogenase</a></li> <li><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a></li> <li><a href="/wiki/Carnitine_palmitoyltransferase_II" title="Carnitine palmitoyltransferase II">Carnitine palmitoyltransferase II</a></li> <li><a href="/wiki/Uncoupling_protein" title="Uncoupling protein">Uncoupling protein</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/Mitochondrial_matrix" title="Mitochondrial matrix">Matrix</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%"><span class="nobold"><i><a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</a></i></span></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/Citrate_synthase" title="Citrate synthase">Citrate synthase</a></li> <li><a href="/wiki/Aconitase" title="Aconitase">Aconitase</a></li> <li><a href="/wiki/Isocitrate_dehydrogenase" title="Isocitrate dehydrogenase">Isocitrate dehydrogenase</a></li> <li><a href="/wiki/Oxoglutarate_dehydrogenase_complex" title="Oxoglutarate dehydrogenase complex">Oxoglutarate dehydrogenase complex</a></li> <li><a href="/wiki/Succinyl_coenzyme_A_synthetase" title="Succinyl coenzyme A synthetase">Succinyl coenzyme A synthetase</a></li> <li><a href="/wiki/Fumarase" title="Fumarase">Fumarase</a></li> <li><a href="/wiki/Malate_dehydrogenase" title="Malate dehydrogenase">Malate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Anaplerotic_reactions" title="Anaplerotic reactions">anaplerotic reactions</a></i></span></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/Aspartate_transaminase" title="Aspartate transaminase">Aspartate transaminase</a></li> <li><a href="/wiki/Glutamate_dehydrogenase" title="Glutamate dehydrogenase">Glutamate dehydrogenase</a></li> <li><a href="/wiki/Pyruvate_dehydrogenase_complex" title="Pyruvate dehydrogenase complex">Pyruvate dehydrogenase complex</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Urea_cycle" title="Urea cycle">urea cycle</a></i></span></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/Carbamoyl_phosphate_synthetase_I" title="Carbamoyl phosphate synthetase I">Carbamoyl phosphate synthetase I</a></li> <li><a href="/wiki/Ornithine_transcarbamylase" title="Ornithine transcarbamylase">Ornithine transcarbamylase</a></li> <li><a href="/wiki/N-Acetylglutamate_synthase" title="N-Acetylglutamate synthase">N-Acetylglutamate synthase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>alcohol metabolism</i></span></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/ALDH2" title="ALDH2">ALDH2</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/PMPCB" title="PMPCB">PMPCB</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/to be sorted</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><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Frataxin" title="Frataxin">Frataxin</a></li> <li><a href="/wiki/Mitochondrial_membrane_transport_protein" title="Mitochondrial membrane transport protein">Mitochondrial membrane transport protein</a> <ul><li><a href="/wiki/Mitochondrial_permeability_transition_pore" title="Mitochondrial permeability transition pore">Mitochondrial permeability transition pore</a></li> <li><a href="/wiki/Mitochondrial_carrier" title="Mitochondrial carrier">Mitochondrial carrier</a></li></ul></li> <li><a href="/wiki/Translocator_protein" title="Translocator protein">Translocator protein</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/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial DNA</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%"><span class="nobold"><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">Complex I</a></span></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/MT-ND1" title="MT-ND1">MT-ND1</a></li> <li><a href="/wiki/MT-ND2" title="MT-ND2">MT-ND2</a></li> <li><a href="/wiki/MT-ND3" title="MT-ND3">MT-ND3</a></li> <li><a href="/wiki/MT-ND4" title="MT-ND4">MT-ND4</a></li> <li><a href="/wiki/MT-ND4L" title="MT-ND4L">MT-ND4L</a></li> <li><a href="/wiki/MT-ND5" title="MT-ND5">MT-ND5</a></li> <li><a href="/wiki/MT-ND6" title="MT-ND6">MT-ND6</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Complex III</a></span></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/MT-CYB" title="MT-CYB">MT-CYB</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Complex IV</a></span></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/Cytochrome_c_oxidase_subunit_I" title="Cytochrome c oxidase subunit I">MT-CO1</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_II" class="mw-redirect" title="Cytochrome c oxidase subunit II">MT-CO2</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_III" title="Cytochrome c oxidase subunit III">MT-CO3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a></span></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/MT-ATP6" title="MT-ATP6">MT-ATP6</a></li> <li><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a class="mw-selflink selflink">tRNA</a></span></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/MT-TA" title="MT-TA">MT-TA</a></li> <li><a href="/wiki/MT-TC" title="MT-TC">MT-TC</a></li> <li><a href="/wiki/MT-TD" title="MT-TD">MT-TD</a></li> <li><a href="/wiki/MT-TE" title="MT-TE">MT-TE</a></li> <li><a href="/wiki/MT-TF" title="MT-TF">MT-TF</a></li> <li><a href="/wiki/MT-TG" title="MT-TG">MT-TG</a></li> <li><a href="/wiki/MT-TH" title="MT-TH">MT-TH</a></li> <li><a href="/wiki/MT-TI" title="MT-TI">MT-TI</a></li> <li><a href="/wiki/MT-TK" title="MT-TK">MT-TK</a></li> <li><a href="/wiki/MT-TL1" title="MT-TL1">MT-TL1</a></li> <li><a href="/wiki/MT-TL2" title="MT-TL2">MT-TL2</a></li> <li><a href="/wiki/MT-TM" title="MT-TM">MT-TM</a></li> <li><a href="/wiki/MT-TN" title="MT-TN">MT-TN</a></li> <li><a href="/wiki/MT-TP" title="MT-TP">MT-TP</a></li> <li><a href="/wiki/MT-TQ" title="MT-TQ">MT-TQ</a></li> <li><a href="/wiki/MT-TR" title="MT-TR">MT-TR</a></li> <li><a href="/wiki/MT-TS1" title="MT-TS1">MT-TS1</a></li> <li><a href="/wiki/MT-TS2" title="MT-TS2">MT-TS2</a></li> <li><a href="/wiki/MT-TT" title="MT-TT">MT-TT</a></li> <li><a href="/wiki/MT-TV_(mitochondrial)" title="MT-TV (mitochondrial)">MT-TV</a></li> <li><a href="/wiki/MT-TW" title="MT-TW">MT-TW</a></li> <li><a href="/wiki/MT-TY" title="MT-TY">MT-TY</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div><i>see also <a href="/wiki/Template:Mitochondrial_diseases" title="Template:Mitochondrial diseases">mitochondrial diseases</a></i></div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐vz5zt Cached time: 20241122140840 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 1.364 seconds Real time usage: 1.631 seconds Preprocessor visited node count: 7256/1000000 Post‐expand include size: 348365/2097152 bytes Template argument size: 3649/2097152 bytes Highest expansion depth: 19/100 Expensive parser function count: 5/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 431326/5000000 bytes Lua time usage: 0.842/10.000 seconds Lua memory usage: 7647375/52428800 bytes Number of Wikibase entities loaded: 1/400 --> <!-- Transclusion expansion time report (%,ms,calls,template) 100.00% 1415.483 1 -total 59.55% 842.988 1 Template:Reflist 44.42% 628.757 89 Template:Cite_journal 10.40% 147.259 8 Template:Navbox 8.47% 119.889 1 Template:Commons_category 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