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vector-toc-landmark"> <div id="vector-toc-pinned-container" class="vector-pinned-container"> <div id="vector-toc" class="vector-toc vector-pinnable-element"> <div class="vector-pinnable-header vector-toc-pinnable-header vector-pinnable-header-pinned" data-feature-name="toc-pinned" data-pinnable-element-id="vector-toc" > <h2 class="vector-pinnable-header-label">目次</h2> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-toc.pin">サイドバーに移動</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">非表示</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">ページ先頭</div> </a> </li> <li id="toc-細胞機構" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#細胞機構"> <div class="vector-toc-text"> 1 細胞機構 </div> </a> <button aria-controls="toc-細胞機構-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> 細胞機構サブセクションを切り替えます </button> <ul id="toc-細胞機構-sublist" class="vector-toc-list"> <li id="toc-miRNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#miRNA"> <div class="vector-toc-text"> 1.1 miRNA </div> </a> <ul id="toc-miRNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-3'_UTRとmiRNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#3'_UTRとmiRNA"> <div class="vector-toc-text"> 1.2 3' UTRとmiRNA </div> </a> <ul id="toc-3'_UTRとmiRNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RISCの活性化と触媒" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RISCの活性化と触媒"> <div class="vector-toc-text"> 1.3 RISCの活性化と触媒 </div> </a> <ul id="toc-RISCの活性化と触媒-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-転写サイレンシング" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#転写サイレンシング"> <div class="vector-toc-text"> 1.4 転写サイレンシング </div> </a> <ul id="toc-転写サイレンシング-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA編集とのクロストーク" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA編集とのクロストーク"> <div class="vector-toc-text"> 1.5 RNA編集とのクロストーク </div> </a> <ul id="toc-RNA編集とのクロストーク-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-生物種間の多様性" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#生物種間の多様性"> <div class="vector-toc-text"> 1.6 生物種間の多様性 </div> </a> <ul id="toc-生物種間の多様性-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-関連する原核生物のシステム" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#関連する原核生物のシステム"> <div class="vector-toc-text"> 1.7 関連する原核生物のシステム </div> </a> <ul id="toc-関連する原核生物のシステム-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-生物学的機能" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#生物学的機能"> <div class="vector-toc-text"> 2 生物学的機能 </div> </a> <button aria-controls="toc-生物学的機能-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> 生物学的機能サブセクションを切り替えます </button> <ul id="toc-生物学的機能-sublist" class="vector-toc-list"> <li id="toc-免疫" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#免疫"> <div class="vector-toc-text"> 2.1 免疫 </div> </a> <ul id="toc-免疫-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-遺伝子のダウンレギュレーション" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#遺伝子のダウンレギュレーション"> <div class="vector-toc-text"> 2.2 遺伝子のダウンレギュレーション </div> </a> <ul id="toc-遺伝子のダウンレギュレーション-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-進化" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#進化"> <div class="vector-toc-text"> 3 進化 </div> </a> <ul id="toc-進化-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-応用" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#応用"> <div class="vector-toc-text"> 4 応用 </div> </a> <button aria-controls="toc-応用-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> 応用サブセクションを切り替えます </button> <ul id="toc-応用-sublist" class="vector-toc-list"> <li id="toc-遺伝子ノックダウンのためのRNAi経路" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#遺伝子ノックダウンのためのRNAi経路"> <div class="vector-toc-text"> 4.1 遺伝子ノックダウンのためのRNAi経路 </div> </a> <ul id="toc-遺伝子ノックダウンのためのRNAi経路-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-機能ゲノミクス" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#機能ゲノミクス"> <div class="vector-toc-text"> 4.2 機能ゲノミクス </div> </a> <ul id="toc-機能ゲノミクス-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-医療" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#医療"> <div class="vector-toc-text"> 4.3 医療 </div> </a> <ul id="toc-医療-sublist" class="vector-toc-list"> <li id="toc-治療応用" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#治療応用"> <div class="vector-toc-text"> 4.3.1 治療応用 </div> </a> <ul id="toc-治療応用-sublist" class="vector-toc-list"> <li id="toc-ウイルス感染" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#ウイルス感染"> <div class="vector-toc-text"> 4.3.1.1 ウイルス感染 </div> </a> <ul id="toc-ウイルス感染-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-がん" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#がん"> <div class="vector-toc-text"> 4.3.1.2 がん </div> </a> <ul id="toc-がん-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-神経疾患" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#神経疾患"> <div class="vector-toc-text"> 4.3.1.3 神経疾患 </div> </a> <ul id="toc-神経疾患-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-治療応用における困難" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#治療応用における困難"> <div class="vector-toc-text"> 4.3.2 治療応用における困難 </div> </a> <ul id="toc-治療応用における困難-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-免疫応答の刺激" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#免疫応答の刺激"> <div class="vector-toc-text"> 4.3.3 免疫応答の刺激 </div> </a> <ul id="toc-免疫応答の刺激-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-治療技術としての展望" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#治療技術としての展望"> <div class="vector-toc-text"> 4.3.4 治療技術としての展望 </div> </a> <ul id="toc-治療技術としての展望-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-バイオテクノロジー" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#バイオテクノロジー"> <div class="vector-toc-text"> 4.4 バイオテクノロジー </div> </a> <ul id="toc-バイオテクノロジー-sublist" class="vector-toc-list"> <li id="toc-食品" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#食品"> <div class="vector-toc-text"> 4.4.1 食品 </div> </a> <ul id="toc-食品-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-その他の作物" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#その他の作物"> <div class="vector-toc-text"> 4.4.2 その他の作物 </div> </a> <ul id="toc-その他の作物-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-殺虫剤" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#殺虫剤"> <div class="vector-toc-text"> 4.4.3 殺虫剤 </div> </a> <ul id="toc-殺虫剤-sublist" class="vector-toc-list"> <li id="toc-トランスジェニック植物" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#トランスジェニック植物"> <div class="vector-toc-text"> 4.4.3.1 トランスジェニック植物 </div> </a> <ul id="toc-トランスジェニック植物-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-外部からの投与" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#外部からの投与"> <div class="vector-toc-text"> 4.4.3.2 外部からの投与 </div> </a> <ul id="toc-外部からの投与-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-ゲノムスケールでのスクリーニング" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#ゲノムスケールでのスクリーニング"> <div class="vector-toc-text"> 4.5 ゲノムスケールでのスクリーニング </div> </a> <ul id="toc-ゲノムスケールでのスクリーニング-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-歴史" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#歴史"> <div class="vector-toc-text"> 5 歴史 </div> </a> <ul id="toc-歴史-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-出典" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#出典"> <div class="vector-toc-text"> 6 出典 </div> </a> <ul id="toc-出典-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-外部リンク" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#外部リンク"> <div class="vector-toc-text"> 7 外部リンク </div> </a> <ul id="toc-外部リンク-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="目次" 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="目次の表示・非表示を切り替え" > <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" > 目次の表示・非表示を切り替え </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">RNA干渉</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="特定の記事の別の言語版に移動します。利用可能な言語45件" > <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-45" aria-hidden="true" > 45の言語版 </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Interferencia_d%27ARN" title="アラゴン語: Interferencia d'ARN" lang="an" hreflang="an" data-title="Interferencia d'ARN" data-language-autonym="Aragonés" data-language-local-name="アラゴン語" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D8%AF%D8%AE%D9%84_%D8%A7%D9%84%D8%B1%D9%86%D8%A7" title="アラビア語: تدخل الرنا" lang="ar" hreflang="ar" data-title="تدخل الرنا" data-language-autonym="العربية" data-language-local-name="アラビア語" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/RNK_interferencija" title="ボスニア語: RNK interferencija" lang="bs" hreflang="bs" data-title="RNK interferencija" data-language-autonym="Bosanski" data-language-local-name="ボスニア語" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/ARN_d%27interfer%C3%A8ncia" title="カタロニア語: ARN d'interferència" lang="ca" hreflang="ca" data-title="ARN d'interferència" data-language-autonym="Català" data-language-local-name="カタロニア語" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/RNA_interference" title="チェコ語: RNA interference" lang="cs" hreflang="cs" data-title="RNA interference" data-language-autonym="Čeština" data-language-local-name="チェコ語" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/RNA-interferens" title="デンマーク語: RNA-interferens" lang="da" hreflang="da" data-title="RNA-interferens" data-language-autonym="Dansk" data-language-local-name="デンマーク語" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/RNA-Interferenz" title="ドイツ語: RNA-Interferenz" lang="de" hreflang="de" data-title="RNA-Interferenz" data-language-autonym="Deutsch" data-language-local-name="ドイツ語" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A0%CE%B1%CF%81%CE%B5%CE%BC%CE%B2%CE%BF%CE%BB%CE%AE_RNA" title="ギリシャ語: Παρεμβολή RNA" lang="el" hreflang="el" data-title="Παρεμβολή RNA" data-language-autonym="Ελληνικά" data-language-local-name="ギリシャ語" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-en badge-Q17437796 badge-featuredarticle mw-list-item" title="秀逸な記事"><a href="https://en.wikipedia.org/wiki/RNA_interference" title="英語: RNA interference" lang="en" hreflang="en" data-title="RNA interference" data-language-autonym="English" data-language-local-name="英語" class="interlanguage-link-target">English</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/ARN_interferente" title="スペイン語: ARN interferente" lang="es" hreflang="es" data-title="ARN interferente" data-language-autonym="Español" data-language-local-name="スペイン語" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/RNA_interferents" title="エストニア語: RNA interferents" lang="et" hreflang="et" data-title="RNA interferents" data-language-autonym="Eesti" data-language-local-name="エストニア語" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Interferentziazko_RNA" title="バスク語: Interferentziazko RNA" lang="eu" hreflang="eu" data-title="Interferentziazko RNA" data-language-autonym="Euskara" data-language-local-name="バスク語" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%D8%AF%D8%A7%D8%AE%D9%84_%D8%A2%D8%B1%D8%A7%D9%86%E2%80%8C%D8%A7%DB%8C" title="ペルシア語: تداخل آران‌ای" lang="fa" hreflang="fa" data-title="تداخل آران‌ای" data-language-autonym="فارسی" data-language-local-name="ペルシア語" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/RNA-interferenssi" title="フィンランド語: RNA-interferenssi" lang="fi" hreflang="fi" data-title="RNA-interferenssi" data-language-autonym="Suomi" data-language-local-name="フィンランド語" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Interf%C3%A9rence_par_ARN" title="フランス語: Interférence par ARN" lang="fr" hreflang="fr" data-title="Interférence par ARN" data-language-autonym="Français" data-language-local-name="フランス語" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/ARN_interferente" title="ガリシア語: ARN interferente" lang="gl" hreflang="gl" data-title="ARN interferente" data-language-autonym="Galego" data-language-local-name="ガリシア語" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/RNAi" title="ヘブライ語: RNAi" lang="he" hreflang="he" data-title="RNAi" data-language-autonym="עברית" data-language-local-name="ヘブライ語" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%86%E0%A4%B0%E0%A4%8F%E0%A4%A8%E0%A4%8F_%E0%A4%B9%E0%A4%B8%E0%A5%8D%E0%A4%A4%E0%A4%95%E0%A5%8D%E0%A4%B7%E0%A5%87%E0%A4%AA" title="ヒンディー語: आरएनए हस्तक्षेप" lang="hi" hreflang="hi" data-title="आरएनए हस्तक्षेप" data-language-autonym="हिन्दी" data-language-local-name="ヒンディー語" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Interferencija_RNK" title="クロアチア語: Interferencija RNK" lang="hr" hreflang="hr" data-title="Interferencija RNK" data-language-autonym="Hrvatski" data-language-local-name="クロアチア語" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/RNS-interferencia" title="ハンガリー語: RNS-interferencia" lang="hu" hreflang="hu" data-title="RNS-interferencia" data-language-autonym="Magyar" data-language-local-name="ハンガリー語" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Interferensi_RNA" title="インドネシア語: Interferensi RNA" lang="id" hreflang="id" data-title="Interferensi RNA" data-language-autonym="Bahasa Indonesia" data-language-local-name="インドネシア語" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/RNA_interference" title="イタリア語: RNA interference" lang="it" hreflang="it" data-title="RNA interference" data-language-autonym="Italiano" data-language-local-name="イタリア語" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%86%E0%B2%B0%E0%B3%8D%E2%80%8C%E0%B2%8E%E0%B2%A8%E0%B3%8D%E2%80%8C%E0%B2%8E_%E0%B2%B9%E0%B2%B8%E0%B3%8D%E0%B2%A4%E0%B2%95%E0%B3%8D%E0%B2%B7%E0%B3%87%E0%B2%AA" title="カンナダ語: ಆರ್‌ಎನ್‌ಎ ಹಸ್ತಕ್ಷೇಪ" lang="kn" hreflang="kn" data-title="ಆರ್‌ಎನ್‌ಎ ಹಸ್ತಕ್ಷೇಪ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="カンナダ語" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/RNA_%EA%B0%84%EC%84%AD" title="韓国語: RNA 간섭" lang="ko" hreflang="ko" data-title="RNA 간섭" data-language-autonym="한국어" data-language-local-name="韓国語" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/RNS_interference" title="ラトビア語: RNS interference" lang="lv" hreflang="lv" data-title="RNS interference" data-language-autonym="Latviešu" data-language-local-name="ラトビア語" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/RNA-interferentie" title="オランダ語: RNA-interferentie" lang="nl" hreflang="nl" data-title="RNA-interferentie" data-language-autonym="Nederlands" data-language-local-name="オランダ語" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/RNA-interferens" title="ノルウェー語(ニーノシュク): RNA-interferens" lang="nn" hreflang="nn" data-title="RNA-interferens" data-language-autonym="Norsk nynorsk" data-language-local-name="ノルウェー語(ニーノシュク)" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/RNA-interferens" title="ノルウェー語(ブークモール): RNA-interferens" lang="nb" hreflang="nb" data-title="RNA-interferens" data-language-autonym="Norsk bokmål" data-language-local-name="ノルウェー語(ブークモール)" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Interferencja_RNA" title="ポーランド語: Interferencja RNA" lang="pl" hreflang="pl" data-title="Interferencja RNA" data-language-autonym="Polski" data-language-local-name="ポーランド語" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AF_RNA_%D9%85%D8%AF%D8%A7%D8%AE%D9%84%D9%87" title="パシュトゥー語: د RNA مداخله" lang="ps" hreflang="ps" data-title="د RNA مداخله" data-language-autonym="پښتو" data-language-local-name="パシュトゥー語" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/ARN_interferente" title="ポルトガル語: ARN interferente" lang="pt" hreflang="pt" data-title="ARN interferente" data-language-autonym="Português" data-language-local-name="ポルトガル語" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ru badge-Q17437798 badge-goodarticle mw-list-item" title="良質な記事"><a href="https://ru.wikipedia.org/wiki/%D0%A0%D0%9D%D0%9A-%D0%B8%D0%BD%D1%82%D0%B5%D1%80%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%8F" title="ロシア語: РНК-интерференция" lang="ru" hreflang="ru" data-title="РНК-интерференция" data-language-autonym="Русский" data-language-local-name="ロシア語" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/RNA_interference" title="スコットランド語: RNA interference" lang="sco" hreflang="sco" data-title="RNA interference" data-language-autonym="Scots" data-language-local-name="スコットランド語" class="interlanguage-link-target">Scots</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/RNA_interference" title="シンプル英語: RNA interference" lang="en-simple" hreflang="en-simple" data-title="RNA interference" data-language-autonym="Simple English" data-language-local-name="シンプル英語" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Interferenca_RNA" title="スロベニア語: Interferenca RNA" lang="sl" hreflang="sl" data-title="Interferenca RNA" data-language-autonym="Slovenščina" data-language-local-name="スロベニア語" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A0%D0%9D%D0%9A_%D0%B8%D0%BD%D1%82%D0%B5%D1%80%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D0%B8%D1%98%D0%B0" title="セルビア語: РНК интерференција" lang="sr" hreflang="sr" data-title="РНК интерференција" data-language-autonym="Српски / srpski" data-language-local-name="セルビア語" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/RNA-interferens" title="スウェーデン語: RNA-interferens" lang="sv" hreflang="sv" data-title="RNA-interferens" data-language-autonym="Svenska" data-language-local-name="スウェーデン語" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%86%E0%AE%B0%E0%AF%8D.%E0%AE%8E%E0%AE%A9%E0%AF%8D.%E0%AE%8F_%E0%AE%95%E0%AF%81%E0%AE%B1%E0%AF%81%E0%AE%95%E0%AF%8D%E0%AE%95%E0%AF%80%E0%AE%9F%E0%AF%81" title="タミル語: ஆர்.என்.ஏ குறுக்கீடு" lang="ta" hreflang="ta" data-title="ஆர்.என்.ஏ குறுக்கீடு" data-language-autonym="தமிழ்" data-language-local-name="タミル語" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/RNA_interference" title="タイ語: RNA interference" lang="th" hreflang="th" data-title="RNA interference" data-language-autonym="ไทย" data-language-local-name="タイ語" class="interlanguage-link-target">ไทย</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/RNA_interferaz" title="トルコ語: RNA interferaz" lang="tr" hreflang="tr" data-title="RNA interferaz" data-language-autonym="Türkçe" data-language-local-name="トルコ語" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A0%D0%9D%D0%9A-%D1%96%D0%BD%D1%82%D0%B5%D1%80%D1%84%D0%B5%D1%80%D0%B5%D0%BD%D1%86%D1%96%D1%8F" title="ウクライナ語: РНК-інтерференція" lang="uk" hreflang="uk" data-title="РНК-інтерференція" data-language-autonym="Українська" data-language-local-name="ウクライナ語" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%A2%D8%B1_%D8%A7%DB%8C%D9%86_%D8%A7%DB%92_%D8%A7%D9%86%D9%B9%D8%B1%D9%81%DB%8C%D8%B1%D9%86%D8%B3" title="ウルドゥー語: آر این اے انٹرفیرنس" lang="ur" hreflang="ur" data-title="آر این اے انٹرفیرنس" data-language-autonym="اردو" data-language-local-name="ウルドゥー語" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Can_thi%E1%BB%87p_RNA" title="ベトナム語: Can thiệp RNA" lang="vi" hreflang="vi" data-title="Can thiệp RNA" data-language-autonym="Tiếng Việt" data-language-local-name="ベトナム語" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/RNA%E5%B9%B2%E6%89%B0" title="中国語: RNA干扰" lang="zh" hreflang="zh" data-title="RNA干扰" data-language-autonym="中文" data-language-local-name="中国語" class="interlanguage-link-target">中文</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/RNA_kan-g%C4%81i" title="閩南語: RNA kan-gāi" lang="nan" hreflang="nan" data-title="RNA kan-gāi" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="閩南語" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li> </ul> <div class="after-portlet after-portlet-lang"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q201993#sitelinks-wikipedia" title="言語間リンクを編集" class="wbc-editpage">リンクを編集</a></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="名前空間"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet 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</div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">出典: フリー百科事典『ウィキペディア（Wikipedia）』</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="ja" dir="ltr"><figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:ShRNA_Lentivirus.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/ShRNA_Lentivirus.svg/330px-ShRNA_Lentivirus.svg.png" decoding="async" width="330" height="419" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/ShRNA_Lentivirus.svg/495px-ShRNA_Lentivirus.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e4/ShRNA_Lentivirus.svg/660px-ShRNA_Lentivirus.svg.png 2x" data-file-width="481" data-file-height="610" /></a><figcaption><a href="/wiki/%E3%83%AC%E3%83%B3%E3%83%81%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9%E5%B1%9E" title="レンチウイルス属">レンチウイルス</a>による哺乳類細胞へのshRNAのデリバリーとRNAi機構</figcaption></figure> RNA干渉（RNAかんしょう、<a href="/wiki/%E8%8B%B1%E8%AA%9E" title="英語">英</a>: RNA interference、RNAi）は、二本鎖<a href="/wiki/%E3%83%AA%E3%83%9C%E6%A0%B8%E9%85%B8" title="リボ核酸">RNA</a>（dsRNA）が<a href="/wiki/%E7%BF%BB%E8%A8%B3_(%E7%94%9F%E7%89%A9%E5%AD%A6)" title="翻訳 (生物学)">翻訳</a>抑制または<a href="/wiki/%E8%BB%A2%E5%86%99_(%E7%94%9F%E7%89%A9%E5%AD%A6)" title="転写 (生物学)">転写</a>抑制によって<a href="/wiki/%E9%81%BA%E4%BC%9D%E5%AD%90%E7%99%BA%E7%8F%BE" title="遺伝子発現">遺伝子の発現</a>を配列特異的に抑制する生物学的過程である。RNAiは歴史的には、"co-suppression"、"post-transcriptional gene silencing"（PTGS）、"quelling"といった名称で知られていた。これらの過程は見かけ上異なるものの、それぞれに対して詳細な研究が行われ、これらの実体はすべてRNAiであることが明らかにされた。<a href="/wiki/%E3%82%A2%E3%83%B3%E3%83%89%E3%83%AA%E3%83%A5%E3%83%BC%E3%83%BB%E3%83%95%E3%82%A1%E3%82%A4%E3%82%A2%E3%83%BC" title="アンドリュー・ファイアー">アンドリュー・ファイアー</a>と<a href="/wiki/%E3%82%AF%E3%83%AC%E3%82%A4%E3%82%B0%E3%83%BB%E3%83%A1%E3%83%AD%E3%83%BC" title="クレイグ・メロー">クレイグ・メロー</a>は、1998年に発表された線虫<a href="/wiki/Caenorhabditis_elegans" class="mw-redirect" title="Caenorhabditis elegans">Caenorhabditis elegans</a>におけるRNAiに関する業績によって、2006年に<a href="/wiki/%E3%83%8E%E3%83%BC%E3%83%99%E3%83%AB%E7%94%9F%E7%90%86%E5%AD%A6%E3%83%BB%E5%8C%BB%E5%AD%A6%E8%B3%9E" title="ノーベル生理学・医学賞">ノーベル生理学・医学賞</a>を受賞した。RNAiとその調節の可能性が発見されて以降、RNAiが目的の遺伝子を抑制する多大な可能性を有していることが明らかとなった。現在ではRNAiは、遺伝子抑制を目的とした<a href="/wiki/%E3%82%A2%E3%83%B3%E3%83%81%E3%82%BB%E3%83%B3%E3%82%B9RNA" title="アンチセンスRNA">アンチセンス</a>治療よりも正確かつ効率的で安定なより良い治療法であることが知られている<a href="#cite_note-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564-1">[1]</a>。 RNAiでは、<a href="/wiki/MiRNA" title="MiRNA">miRNA</a>と<a href="/wiki/SiRNA" title="SiRNA">siRNA</a>という2つのタイプの低分子RNAが中心的役割を果たす。RNAは遺伝子の直接産物であり、こうした低分子RNAは酵素複合体を指揮し、<a href="/wiki/%E4%BC%9D%E4%BB%A4RNA" title="伝令RNA">mRNA</a>の分解や翻訳の阻害といった、転写後段階での遺伝子サイレンシングを介して標的遺伝子の活性を低下させる。さらに、siRNAやmiRNAに相補的なゲノム領域でDNAのメチル化を触媒する酵素複合体による、転写前段階でのサイレンシング機構も存在し、それによって転写が阻害されることもある。RNAiは、<a href="/wiki/%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ウイルス">ウイルス</a>や<a href="/wiki/%E3%83%88%E3%83%A9%E3%83%B3%E3%82%B9%E3%83%9D%E3%82%BE%E3%83%B3" title="トランスポゾン">トランスポゾン</a>といった寄生性のヌクレオチド配列に対する細胞の防御に重要な役割を果たす。また、発生にも影響を与える。 RNAi経路は<a href="/wiki/%E5%8B%95%E7%89%A9" title="動物">動物</a>を含む多くの<a href="/wiki/%E7%9C%9F%E6%A0%B8%E7%94%9F%E7%89%A9" title="真核生物">真核生物</a>でみられ、<a href="/wiki/Dicer" title="Dicer">Dicer</a>によって開始される。Dicerは長いdsRNA分子を約21<a href="/wiki/%E3%83%8C%E3%82%AF%E3%83%AC%E3%82%AA%E3%83%81%E3%83%89" title="ヌクレオチド">ヌクレオチド</a>からなる短いsiRNA二本鎖断片へと切断する酵素である。その後、siRNAは2つの一本鎖RNA（ssRNA）、すなわちパッセンジャー鎖とガイド鎖へと巻き戻される。パッセンジャー鎖は分解され、ガイド鎖は<a href="/wiki/RNA%E8%AA%98%E5%B0%8E%E3%82%B5%E3%82%A4%E3%83%AC%E3%83%B3%E3%82%B7%E3%83%B3%E3%82%B0%E8%A4%87%E5%90%88%E4%BD%93" title="RNA誘導サイレンシング複合体">RNA誘導サイレンシング複合体</a>（RISC）へと取り込まれる。RNAi経路の中で最もよく研究されているのは転写後段階での遺伝子サイレンシングであり、この過程ではガイド鎖がmRNA分子中の相補的な配列と対合し、RISCの触媒要素である<a href="/wiki/%E3%82%A2%E3%83%AB%E3%82%B4%E3%83%8E%E3%83%BC%E3%83%88_(%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA)" title="アルゴノート (タンパク質)">Argonaute</a>2（Ago2）による切断が誘導される。一部の生物種では、当初のsiRNA濃度が限定的なものであっても、この過程が増幅し、全身に拡散する。 細胞に導入された合成dsRNAは目的の遺伝子の選択的かつ強固な抑制を誘導することとができるため、RNAiは培養細胞においても個体レベルにおいても有益な研究ツールである。RNAiは細胞内の各遺伝子を体系的にオフにする大規模スクリーニングに利用することができ、<a href="/wiki/%E7%B4%B0%E8%83%9E%E5%88%86%E8%A3%82" title="細胞分裂">細胞分裂</a>などのイベントや特定の細胞過程に必要な要素の同定のために活用することができる。また、この経路は<a href="/wiki/%E3%83%90%E3%82%A4%E3%82%AA%E3%83%86%E3%82%AF%E3%83%8E%E3%83%AD%E3%82%B8%E3%83%BC" class="mw-redirect" title="バイオテクノロジー">バイオテクノロジー</a>や<a href="/wiki/%E5%8C%BB%E7%99%82" title="医療">医療</a>、そして<a href="/wiki/%E6%AE%BA%E8%99%AB%E5%89%A4" title="殺虫剤">殺虫剤</a>など実用的なツールとしても利用されている<a href="#cite_note-s1308-2">[2]</a>。 <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="細胞機構">細胞機構</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=1" title="節を編集: 細胞機構">編集</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:2ffl-by-domain.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/2ffl-by-domain.png/220px-2ffl-by-domain.png" decoding="async" width="220" height="371" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/2ffl-by-domain.png/330px-2ffl-by-domain.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/2ffl-by-domain.png/440px-2ffl-by-domain.png 2x" data-file-width="690" data-file-height="1164" /></a><figcaption><a href="/wiki/%E3%82%B8%E3%82%A2%E3%83%AB%E3%82%B8%E3%82%A2" title="ジアルジア">ジアルジア</a>Giardia intestinalis由来Dicerタンパク質。dsRNAからsiRNAへの切断を触媒する。RNaseドメインは緑色、PAZドメインは黄色、プラットホームドメインは赤色、コネクターヘリックスは青色で示されている<a href="#cite_note-Macrae-3">[3]</a>。</figcaption></figure> RNAiは、RNA誘導サイレンシング複合体（RISC）によって制御されるRNA依存的遺伝子サイレンシング過程であり、<a href="/wiki/%E7%B4%B0%E8%83%9E%E8%B3%AA" title="細胞質">細胞質</a>に存在する短い二本鎖RNA（dsRNA）分子によって開始される。dsRNAが外因性のもの（RNAゲノムを持つ<a href="/wiki/%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ウイルス">ウイルス</a>の感染や実験室的操作に由来するもの）である場合には、RNAは直接細胞質に取り込まれ、Dicerによって短い断片へと切断される。ゲノム中のRNAコーディング遺伝子から発現したpre-miRNAなどのように、RNAi経路を開始するdsRNAは内因性のもの（細胞に由来するもの）である場合もある。こうした遺伝子に由来する<a href="/wiki/%E4%B8%80%E6%AC%A1%E8%BB%A2%E5%86%99%E7%94%A3%E7%89%A9" title="一次転写産物">一次転写産物</a>は、まず<a href="/wiki/%E7%B4%B0%E8%83%9E%E6%A0%B8" title="細胞核">核</a>内でpre-miRNAに特徴的な<a href="/wiki/%E3%82%B9%E3%83%86%E3%83%A0%E3%83%AB%E3%83%BC%E3%83%97" title="ステムループ">ステムループ</a>構造を形成するようプロセシングされる。その後、外因性と内因性の2つのdsRNA経路はRISCへと集約される<a href="#cite_note-pmid15614608-4">[4]</a>。 外因性のdsRNAは<a href="/wiki/%E3%83%AA%E3%83%9C%E3%83%8C%E3%82%AF%E3%83%AC%E3%82%A2%E3%83%BC%E3%82%BC" title="リボヌクレアーゼ">リボヌクレアーゼ</a>タンパク質であるDicerによってRNAiを開始する<a href="#cite_note-Bernstein-5">[5]</a>。Dicerは植物ではdsRNA、ヒトでは<a href="/wiki/ShRNA" title="ShRNA">shRNA</a>と結合して切断を行い、3'末端に2ヌクレオチドの突出部を持つ20–25塩基対の二本鎖断片を形成する<a href="#cite_note-:2-6">[6]</a>。この長さは、標的遺伝子に対する特異性を最大化し、かつ非特異的な効果を最小化することが複数の生物種のゲノムに対する<a href="/wiki/%E3%83%90%E3%82%A4%E3%82%AA%E3%82%A4%E3%83%B3%E3%83%95%E3%82%A9%E3%83%9E%E3%83%86%E3%82%A3%E3%82%AF%E3%82%B9" title="バイオインフォマティクス">バイオインフォマティクス</a>研究から示唆されている<a href="#cite_note-Qiu-7">[7]</a>。こうした短い二本鎖断片はsiRNAと呼ばれる。その後、RISCローディング複合体（RLC）によって、siRNAは一本鎖へと分離されて活性型のRISCへ取り込まれる。<a href="/wiki/%E3%82%B7%E3%83%A7%E3%82%A6%E3%82%B8%E3%83%A7%E3%82%A6%E3%83%90%E3%82%A8" title="ショウジョウバエ">ショウジョウバエ</a>のRLCはDcr-2（Dicer2）とR2D2を含み、Ago2とRISCの一体化のために重要である<a href="#cite_note-Nakanishi-8">[8]</a>。<a href="/w/index.php?title=TAF11&action=edit&redlink=1" class="new" title="「TAF11」 (存在しないページ)">TAF11</a>（<a href="https://en.wikipedia.org/wiki/TAF11" class="extiw" title="en:TAF11">英語版</a>）はDcr-2とR2D2の四量体化を促進してRLCを組み立て、siRNAに対する結合親和性を10倍増加させる。TAF11との結合によって、R2D2/Dcr2-initiator（RDI）複合体はRLCへと変換される<a href="#cite_note-:3-9">[9]</a>。R2D2にはタンデムに並んだdsRNA結合ドメインが存在し、siRNA二本鎖の熱力学的に安定な末端を認識する。一方、Dcr-2は熱力学的安定性の低い末端を認識する。RISCへのRNAのローディングは非対称的であり、Ago2のMIDドメインはsiRNAの熱力学的に不安定な末端を認識する。そのため、5'末端がMIDドメインに認識されなかったパッセンジャー鎖は放出され、もう一方のガイド鎖はAgoと協調的にRISCを形成する<a href="#cite_note-Nakanishi-8">[8]</a>。 RISCへと取り込まれた後、siRNAは標的mRNAと塩基対を形成して切断を行い、そのmRNAが翻訳の鋳型として利用されることを防ぐ<a href="#cite_note-:4-10">[10]</a>。siRNAとは異なり、miRNAがロードされたRISCはmRNA上の相補性領域を探してスキャンする。 miRNAは通常不完全な相補性でmRNAの<a href="/wiki/3%27_%E9%9D%9E%E7%BF%BB%E8%A8%B3%E9%A0%98%E5%9F%9F" title="3' 非翻訳領域">3' UTR</a>領域に結合し、<a href="/wiki/%E3%83%AA%E3%83%9C%E3%82%BD%E3%83%BC%E3%83%A0" title="リボソーム">リボソーム</a>が翻訳のためにアクセスすることを防ぐ役割を果たす<a href="#cite_note-:5-11">[11]</a>。 <div class="mw-heading mw-heading3"><h3 id="miRNA">miRNA</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=2" title="節を編集: miRNA">編集</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Microrna_secondary_structure.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/220px-Microrna_secondary_structure.png" decoding="async" width="220" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/330px-Microrna_secondary_structure.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/440px-Microrna_secondary_structure.png 2x" data-file-width="684" data-file-height="429" /></a><figcaption><a href="/wiki/%E3%83%A4%E3%82%BB%E3%82%A4%E3%82%AB%E3%83%B3%E3%83%A9%E3%83%B3" title="ヤセイカンラン">Brassica oleracea</a>由来pri-miRNAのステムループ型<a href="/wiki/%E6%A0%B8%E9%85%B8%E3%81%AE%E4%BA%8C%E6%AC%A1%E6%A7%8B%E9%80%A0" title="核酸の二次構造">二次構造</a></figcaption></figure> <div class="rellink" style="margin-bottom: 0.5em; padding-left: 2em; font-size: 90%;" role="note">→詳細は「<a href="/wiki/MiRNA" title="MiRNA">miRNA</a>」を参照</div> miRNAは<a href="/wiki/%E3%82%B2%E3%83%8E%E3%83%A0" title="ゲノム">ゲノム</a>にコードされた<a href="/wiki/%E3%83%8E%E3%83%B3%E3%82%B3%E3%83%BC%E3%83%87%E3%82%A3%E3%83%B3%E3%82%B0RNA" title="ノンコーディングRNA">ノンコーディングRNA</a>であり、特に発生過程において遺伝子発現の調節を補助する<a href="#cite_note-Zhao_Y_2007-12">[12]</a>。広義のRNAiには、外来dsRNAから産生されるsiRNAによるものに加え、miRNAによる内因性の遺伝子サイレンシング効果も含まれる。成熟したmiRNAは外因性dsRNAから産生されたsiRNAと構造的に類似しているが、成熟するまでに広範囲の<a href="/wiki/%E8%BB%A2%E5%86%99%E5%BE%8C%E4%BF%AE%E9%A3%BE" title="転写後修飾">転写後修飾</a>を必要とする。miRNAは最終産物よりもずっと長いRNAコーディング遺伝子からpri-miRNA（primary miRNA）と呼ばれる一次転写産物として発現し、核内で<a href="/w/index.php?title=%E3%83%9E%E3%82%A4%E3%82%AF%E3%83%AD%E3%83%97%E3%83%AD%E3%82%BB%E3%83%83%E3%82%B5%E3%83%BC%E8%A4%87%E5%90%88%E4%BD%93&action=edit&redlink=1" class="new" title="「マイクロプロセッサー複合体」 (存在しないページ)">マイクロプロセッサー複合体</a>（<a href="https://en.wikipedia.org/wiki/Microprocessor_complex" class="extiw" title="en:Microprocessor complex">英語版</a>）によってステムループ構造を持つ約70ヌクレオチドの長さのpre-miRNA（precursor miRNA）へとプロセシングされる。マイクロプロセッサー複合体には、<a href="/w/index.php?title=Drosha&action=edit&redlink=1" class="new" title="「Drosha」 (存在しないページ)">Drosha</a>（<a href="https://en.wikipedia.org/wiki/Drosha" class="extiw" title="en:Drosha">英語版</a>）と呼ばれる<a href="/w/index.php?title=%E3%83%AA%E3%83%9C%E3%83%8C%E3%82%AF%E3%83%AC%E3%82%A2%E3%83%BC%E3%82%BCIII&action=edit&redlink=1" class="new" title="「リボヌクレアーゼIII」 (存在しないページ)">RNase III</a>（<a href="https://en.wikipedia.org/wiki/Ribonuclease_III" class="extiw" title="en:Ribonuclease III">英語版</a>）酵素とdsRNA結合タンパク質<a href="/w/index.php?title=DGCR8&action=edit&redlink=1" class="new" title="「DGCR8」 (存在しないページ)">DGCR8</a>（<a href="https://en.wikipedia.org/wiki/Microprocessor_complex_subunit_DGCR8" class="extiw" title="en:Microprocessor complex subunit DGCR8">英語版</a>）が含まれる。Dicerはこのpre-miRNAのdsRNA部分に結合して切断を行い、RISCに取り込まれる成熟型miRNA分子が産生される。これより下流では、miRNAとsiRNAは同様の装置を利用する<a href="#cite_note-Denli-13">[13]</a>。ウイルス由来のmiRNAとしては、<a href="/wiki/%E3%82%A8%E3%83%97%E3%82%B9%E3%82%BF%E3%82%A4%E3%83%B3%E3%83%BB%E3%83%90%E3%83%BC%E3%83%AB%E3%83%BB%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="エプスタイン・バール・ウイルス">エプスタイン・バール・ウイルス</a>（EBV）にコードされたものが最初に記載された<a href="#cite_note-Pfeffer_734–736-14">[14]</a>。それ以降、ウイルスでは多くのmiRNAの記載がなされている。VIRmiRNAは、ウイルス性のmiRNAとその標的、そして抗ウイルス性miRNAに関する総合的なカタログである<a href="#cite_note-Qureshi_bau103-15">[15]</a>。 miRNAは長いdsRNA前駆体に由来するsiRNAとはいくつかの点で異なる。特に動物では、miRNAと標的mRNAとの塩基対形成は不完全であることが一般的であり、また類似した配列を持つ多くの異なるmRNAの翻訳を阻害する。対照的に、siRNAは通常は完全な塩基対形成を行い、唯一の特異的な標的に対してのみmRNAの切断を誘導する<a href="#cite_note-Pillai_2006-16">[16]</a>。ショウジョウバエやC. elegansでは、miRNAとsiRNAはそれぞれ異なるArgonauteタンパク質とDicer酵素によってプロセシングされる<a href="#cite_note-Okamura-17">[17]</a><a href="#cite_note-Lee-18">[18]</a>。 <div class="mw-heading mw-heading3"><h3 id="3'_UTRとmiRNA">3' UTRとmiRNA</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=3" title="節を編集: 3' UTRとmiRNA">編集</a>]</div> <div class="rellink" style="margin-bottom: 0.5em; padding-left: 2em; font-size: 90%;" role="note">→詳細は「<a href="/wiki/3%27_%E9%9D%9E%E7%BF%BB%E8%A8%B3%E9%A0%98%E5%9F%9F" title="3' 非翻訳領域">3' 非翻訳領域</a>」を参照</div> mRNAの3' UTRには、転写後にRNAiを引き起こす調節配列が存在することが多い。こうした3' UTRには、miRNAの結合部位と調節タンパク質の結合部位の双方が存在することが多い。miRNAは3' UTR内の特定の部位に結合することで、翻訳の阻害または転写産物の分解によって遺伝子発現を低下させる。また、3' UTRにはmRNAの発現を阻害する<a href="/wiki/%E3%83%AA%E3%83%97%E3%83%AC%E3%83%83%E3%82%B5%E3%83%BC" title="リプレッサー">リプレッサー</a>タンパク質が結合する<a href="/wiki/%E3%82%B5%E3%82%A4%E3%83%AC%E3%83%B3%E3%82%B5%E3%83%BC_(%E9%81%BA%E4%BC%9D%E5%AD%A6)" title="サイレンサー (遺伝学)">サイレンサー</a>領域が存在する場合もある。 miRNAの配列とアノテーションがアーカイブされているウェブサイト<a href="/w/index.php?title=MiRBase&action=edit&redlink=1" class="new" title="「MiRBase」 (存在しないページ)">miRBase</a>（<a href="https://en.wikipedia.org/wiki/miRBase" class="extiw" title="en:miRBase">英語版</a>）<a href="#cite_note-19">[19]</a>には、2014年時点で233の生物種の28,645種類のエントリが登録されている。miRNAには平均して約400種類の標的ｍRNAが存在する（数百の遺伝子の発現に影響を与える）ことが予測されている<a href="#cite_note-Friedman-20">[20]</a>。ヒトのmRNAの3' UTRにはバックグラウンドレベルよりも高い水準で保存されている標的部位が45,000か所以上存在し、タンパク質コーディング遺伝子の60%以上に対してmiRNAとの対合を維持するような選択圧がはたらいていると推定されている<a href="#cite_note-Friedman-20">[20]</a>。 1種類のmiRNAが数百種類のmRNAの安定性を低下させる場合があることは、直接的な実験により示されている<a href="#cite_note-pmid15685193-21">[21]</a>。一方他の実験では、1種類のmiRNAが数百種類のタンパク質の産生を抑制する可能性があるものの、多くの場合こうした抑制は比較的弱いもの（2倍未満）であることが示されている<a href="#cite_note-:6-22">[22]</a><a href="#cite_note-:7-23">[23]</a>。 miRNAによる遺伝子発現の調節の異常は、<a href="/wiki/%E6%82%AA%E6%80%A7%E8%85%AB%E7%98%8D" title="悪性腫瘍">がん</a>において重要であるようである<a href="#cite_note-pmid21931505-24">[24]</a>。一例として、消化器がんでは9種類のmiRNAに<a href="/wiki/%E3%82%A8%E3%83%94%E3%82%B8%E3%82%A7%E3%83%8D%E3%83%86%E3%82%A3%E3%82%AF%E3%82%B9" title="エピジェネティクス">エピジェネティック</a>な変化が生じ、<a href="/wiki/DNA%E4%BF%AE%E5%BE%A9" title="DNA修復">DNA修復</a>酵素をダウンレギュレーションする作用を示していることが同定されている<a href="#cite_note-pmid25987950-25">[25]</a>。 miRNAによる遺伝子発現調節の異常は、<a href="/wiki/%E7%B5%B1%E5%90%88%E5%A4%B1%E8%AA%BF%E7%97%87" title="統合失調症">統合失調症</a>、<a href="/wiki/%E5%8F%8C%E6%A5%B5%E6%80%A7%E9%9A%9C%E5%AE%B3" title="双極性障害">双極性障害</a>、<a href="/wiki/%E5%A4%A7%E3%81%86%E3%81%A4%E7%97%85" class="mw-redirect" title="大うつ病">大うつ病</a>、<a href="/wiki/%E3%83%91%E3%83%BC%E3%82%AD%E3%83%B3%E3%82%BD%E3%83%B3%E7%97%85" title="パーキンソン病">パーキンソン病</a>、<a href="/wiki/%E3%82%A2%E3%83%AB%E3%83%84%E3%83%8F%E3%82%A4%E3%83%9E%E3%83%BC%E7%97%85" title="アルツハイマー病">アルツハイマー病</a>、<a href="/wiki/%E8%87%AA%E9%96%89%E7%97%87%E3%82%B9%E3%83%9A%E3%82%AF%E3%83%88%E3%83%A9%E3%83%A0%E9%9A%9C%E5%AE%B3" class="mw-redirect" title="自閉症スペクトラム障害">自閉症スペクトラム障害</a>など精神神経疾患にも重要であるようである<a href="#cite_note-pmid24653674-26">[26]</a><a href="#cite_note-pmid22539927-27">[27]</a><a href="#cite_note-pmid25636176-28">[28]</a>。 <div class="mw-heading mw-heading3"><h3 id="RISCの活性化と触媒">RISCの活性化と触媒</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=4" title="節を編集: RISCの活性化と触媒">編集</a>]</div> 外因性のdsRNAには、C. elegansではRDE-4、ショウジョウバエではR2D2と呼ばれるエフェクタータンパク質が検知して結合し、Dicerの活性を刺激する<a href="#cite_note-Parker-29">[29]</a>。このタンパク質は長いdsRNAにのみ結合するが、こうした長さに対する特異性を生み出す機構は不明である<a href="#cite_note-Parker-29">[29]</a>。その後、このRNA結合タンパク質は切断されたsiRNAのRISCへの移行を促進する<a href="#cite_note-Liu2003-30">[30]</a>。 C. elegansでは、Dicerによって産生された「一次性」のsiRNAを鋳型として「二次性」のsiRNAが合成されることで、この開始応答は増幅される<a href="#cite_note-Baulcombe_D_2007_199–200-31">[31]</a>。こうした「二次性」のsiRNAはDicerによって産生されたsiRNAとは構造的に異なり、<a href="/wiki/RNA%E4%BE%9D%E5%AD%98%E6%80%A7RNA%E3%83%9D%E3%83%AA%E3%83%A1%E3%83%A9%E3%83%BC%E3%82%BC" title="RNA依存性RNAポリメラーゼ">RNA依存性RNAポリメラーゼ</a>（RdRP）によって産生されているようである<a href="#cite_note-Pak-32">[32]</a><a href="#cite_note-Sijen-33">[33]</a>。<figure typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:RNAi_overview.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/RNAi_overview.png/400px-RNAi_overview.png" decoding="async" width="400" height="237" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/RNAi_overview.png/600px-RNAi_overview.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0a/RNAi_overview.png/800px-RNAi_overview.png 2x" data-file-width="7289" data-file-height="4319" /></a><figcaption>低分子RNAの生合成: pri-miRNAは核内で転写された後、ヘアピン型に折り返され、マイクロプロセッサー複合体によってトリミングされて60–70ヌクレオチドのpre-miRNAが形成される。pre-miRNAは<a href="/wiki/%E6%A0%B8%E8%86%9C%E5%AD%94%E8%A4%87%E5%90%88%E4%BD%93" class="mw-redirect" title="核膜孔複合体">核膜孔複合体</a>（NPC）を通って細胞質へ輸送され、そこでDicerによって約20ヌクレオチドのmiRNA二本鎖へとさらにトリミングされる（pre-siRNAもこの段階で経路に加わる）。その後、この二本鎖はAgoへロードされてpre-RISCが形成され、パッセンジャー鎖が放出されて活性型RISCが形成される。</figcaption></figure> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Argonaute_1u04_1ytu_composite.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/Argonaute_1u04_1ytu_composite.png/300px-Argonaute_1u04_1ytu_composite.png" decoding="async" width="300" height="121" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/Argonaute_1u04_1ytu_composite.png/450px-Argonaute_1u04_1ytu_composite.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/90/Argonaute_1u04_1ytu_composite.png/600px-Argonaute_1u04_1ytu_composite.png 2x" data-file-width="2760" data-file-height="1110" /></a><figcaption>左: <a href="/wiki/%E5%8F%A4%E7%B4%B0%E8%8F%8C" title="古細菌">古細菌</a> <a href="/wiki/Pyrococcus_furiosus" class="mw-redirect" title="Pyrococcus furiosus">Pyrococcus furiosus</a>由来のArgonauteタンパク質全長構造。右: dsRNAと複合体を形成したArgonauteタンパク質の<a href="/w/index.php?title=Piwi&action=edit&redlink=1" class="new" title="「Piwi」 (存在しないページ)">Piwi</a>（<a href="https://en.wikipedia.org/wiki/Piwi" class="extiw" title="en:Piwi">英語版</a>）ドメイン。</figcaption></figure> RISCの活性を担う構成要素はArgonauteタンパク質と呼ばれる<a href="/wiki/%E3%82%A8%E3%83%B3%E3%83%89%E3%83%8C%E3%82%AF%E3%83%AC%E3%82%A2%E3%83%BC%E3%82%BC" title="エンドヌクレアーゼ">エンドヌクレアーゼ</a>であり、結合したsiRNAに相補的な標的mRNAを切断する<a href="#cite_note-Daneholt2006-34">[34]</a>。Dicerによって形成される断片は二本鎖であるため、理論上は双方の鎖が機能的なsiRNAとなることができる。しかしながら、2本の鎖のうちArgonauteタンパク質に結合し、遺伝子サイレンシングを指揮するのは、ガイド鎖と呼ばれる一方の鎖のみである。もう一方の鎖はパッセンジャー鎖またはアンチガイド鎖と呼ばれ、RISCの活性化の過程で分解される<a href="#cite_note-Gregory-35">[35]</a>。当初は<a href="/wiki/%E3%82%A2%E3%83%87%E3%83%8E%E3%82%B7%E3%83%B3%E4%B8%89%E3%83%AA%E3%83%B3%E9%85%B8" title="アデノシン三リン酸">ATP</a>依存的な<a href="/wiki/%E3%83%98%E3%83%AA%E3%82%AB%E3%83%BC%E3%82%BC" title="ヘリカーゼ">ヘリカーゼ</a>によって2つの鎖が分離されると考えられていたが<a href="#cite_note-Lodish-36">[36]</a>、この過程は実際にはATP非依存的であり、RISCの構成要素によって直接行われることが示されている<a href="#cite_note-Matranga-37">[37]</a><a href="#cite_note-Leuschner-38">[38]</a>。しかしながら、in vitroにおけるATP存在下と非存在下でのRNAiの速度論的解析からは、触媒後の複合体から切断されたmRNAを巻き戻して除去するためにATPが必要である可能性が示されている<a href="#cite_note-Haley_2004_599–606-39">[39]</a>。ガイド鎖は5'末端の対合の安定性が低い傾向があるが<a href="#cite_note-:8-40">[40]</a>、RISCへの取り込みの前のDicerによるdsRNAの切断の方向には影響を与えない<a href="#cite_note-Preall-41">[41]</a>。Dicerではなく、R2D2タンパク質がより安定性の高いパッセンジャー鎖の5'末端に結合することが鎖の識別因子として機能している可能性がある<a href="#cite_note-Tomari-42">[42]</a>。 Argonauteタンパク質のRNA結合の構造的基盤は、RNAが結合したArgonauteタンパク質のドメインの<a href="/wiki/X%E7%B7%9A%E7%B5%90%E6%99%B6%E6%A7%8B%E9%80%A0%E8%A7%A3%E6%9E%90" title="X線結晶構造解析">X線結晶構造解析</a>によって研究されている。RNA鎖のリン酸化5'末端はArgonauteタンパク質の保存された塩基性ポケットに入り、<a href="/wiki/%E3%83%9E%E3%82%B0%E3%83%8D%E3%82%B7%E3%82%A6%E3%83%A0" title="マグネシウム">マグネシウム</a>などの二価<a href="/wiki/%E3%82%AB%E3%83%81%E3%82%AA%E3%83%B3" class="mw-redirect" title="カチオン">カチオン</a>を介して、また5'末端のヌクレオチドと保存された<a href="/wiki/%E3%83%81%E3%83%AD%E3%82%B7%E3%83%B3" title="チロシン">チロシン</a>残基との<a href="/wiki/%CE%A0-%CF%80%E7%9B%B8%E4%BA%92%E4%BD%9C%E7%94%A8" title="Π-π相互作用">スタッキング</a>によって接触を行っている。この部位はsiRNAが標的mRNAへ結合する際の核形成部位となると考えられている<a href="#cite_note-Ma-43">[43]</a>。ガイド鎖の5'末端または3'末端のミスマッチによる阻害効果の解析からは、ガイド鎖の5'末端は標的mRNAとのマッチングと結合を担う一方で、3'末端は標的mRNAをRISCが切断を行いやすい領域へ物理的に配置をする役割を担っている可能性が高いことが示されている<a href="#cite_note-Haley_2004_599–606-39">[39]</a>。 活性化されたRISC複合体が細胞内のmRNA標的をどのように見つけているのかに関しては未解明である。切断過程と翻訳との関連性が提唱されているが、標的mRNAの翻訳はRNAiを介した分解に必要不可欠なものではない<a href="#cite_note-:9-44">[44]</a>。事実、RNAiは翻訳されていない標的mRNAに対してより効率的に働いている可能性がある<a href="#cite_note-Gu-45">[45]</a>。Argonauteタンパク質は<a href="/w/index.php?title=P-body&action=edit&redlink=1" class="new" title="「P-body」 (存在しないページ)">P-body</a>（<a href="https://en.wikipedia.org/wiki/P-bodies" class="extiw" title="en:P-bodies">英語版</a>）（cytoplasmic body、GW bodyとも）と呼ばれる細胞質の特定の領域に局在している。この領域ではｍRNAの分解が高率で行われており<a href="#cite_note-SenBlau-46">[46]</a>、miRNAの活性もP-bodyに集中している<a href="#cite_note-Lian-47">[47]</a>。P-bodyの破壊によってRNAiの効率が低下することから、この部位がRNAi過程に重要であることが示唆される<a href="#cite_note-Jakymiw-48">[48]</a>。 <div class="mw-heading mw-heading3"><h3 id="転写サイレンシング">転写サイレンシング</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=5" title="節を編集: 転写サイレンシング">編集</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:RNAi-simplified.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/RNAi-simplified.png/350px-RNAi-simplified.png" decoding="async" width="350" height="299" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/RNAi-simplified.png/525px-RNAi-simplified.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/RNAi-simplified.png/700px-RNAi-simplified.png 2x" data-file-width="800" data-file-height="684" /></a><figcaption>DicerはdsRNAをトリミングし、siRNAまたはmiRNAを形成する。こうしたプロセシングを受けたRNAはRISCに取り込まれ、DNAや新生RNA鎖を標的として転写をサイレンシングしたり、mRNAを標的として翻訳を阻害したりする<a href="#cite_note-Hammond2000-49">[49]</a>。</figcaption></figure> RNAi経路の構成要素は、多くの真核生物においてゲノムの組織化と構造の維持に利用されている。<a href="/wiki/%E3%83%92%E3%82%B9%E3%83%88%E3%83%B3" title="ヒストン">ヒストン</a>の修飾とそれに関連した<a href="/wiki/%E3%83%98%E3%83%86%E3%83%AD%E3%82%AF%E3%83%AD%E3%83%9E%E3%83%81%E3%83%B3" title="ヘテロクロマチン">ヘテロクロマチン</a>形成の誘導は、転写前段階で遺伝子をダウンレギュレーションする役割を果たす<a href="#cite_note-Holmquist_2006-50">[50]</a>。この過程は<a href="/w/index.php?title=RNA%E8%AA%98%E5%B0%8E%E8%BB%A2%E5%86%99%E3%82%B5%E3%82%A4%E3%83%AC%E3%83%B3%E3%82%B7%E3%83%B3%E3%82%B0&action=edit&redlink=1" class="new" title="「RNA誘導転写サイレンシング」 (存在しないページ)">RNA誘導転写サイレンシング</a>（<a href="https://en.wikipedia.org/wiki/RNA-induced_transcriptional_silencing" class="extiw" title="en:RNA-induced transcriptional silencing">英語版</a>）（RITS）と呼ばれ、RITS複合体と呼ばれるタンパク質複合体によって行われる。<a href="/wiki/%E5%88%86%E8%A3%82%E9%85%B5%E6%AF%8D" title="分裂酵母">分裂酵母</a>Schizosaccharomyces pombeでは、この複合体にはArgonaute、<a href="/wiki/%E3%82%AF%E3%83%AD%E3%83%A2%E3%83%89%E3%83%A1%E3%82%A4%E3%83%B3" title="クロモドメイン">クロモドメイン</a>タンパク質Chp1、そしてTas3と呼ばれる機能未知タンパク質が含まれる<a href="#cite_note-Verdel-51">[51]</a>。ヘテロクロマチン領域の誘導と拡大にはArgonauteとRdRPタンパク質が必要である<a href="#cite_note-Irvine-52">[52]</a>。これらの遺伝子を欠失させた分裂酵母は、<a href="/w/index.php?title=%E3%83%92%E3%82%B9%E3%83%88%E3%83%B3%E3%83%A1%E3%83%81%E3%83%AB%E5%8C%96&action=edit&redlink=1" class="new" title="「ヒストンメチル化」 (存在しないページ)">ヒストンメチル化</a>（<a href="https://en.wikipedia.org/wiki/Histone_methylation" class="extiw" title="en:Histone methylation">英語版</a>）と<a href="/wiki/%E3%82%BB%E3%83%B3%E3%83%88%E3%83%AD%E3%83%A1%E3%82%A2" title="セントロメア">セントロメア</a>形成が破壊され<a href="#cite_note-Volpe_2002-53">[53]</a>、<a href="/wiki/%E7%B4%B0%E8%83%9E%E5%88%86%E8%A3%82" title="細胞分裂">細胞分裂</a>の進行は遅くなるか、もしくは<a href="/wiki/%E5%BE%8C%E6%9C%9F_(%E7%B4%B0%E8%83%9E%E5%88%86%E8%A3%82)" title="後期 (細胞分裂)">後期</a>の段階で停止する<a href="#cite_note-Volpe_2003-54">[54]</a>。またある場合には、ヒストン修飾と関係した同様の過程によって遺伝子の転写がアップレギュレーションされることも観察されている<a href="#cite_note-Li-55">[55]</a>。 RITS複合体がヘテロクロマチンの形成や組織化を誘導する機構の詳細は未解明である。多くの研究は分裂酵母の接合型遺伝子座に焦点を当てているが、この遺伝子座における活性は他の生物やゲノム領域における活性を代表するものではない可能性もある。既存のヘテロクロマチン領域の維持に際しては、RITSはその領域の遺伝子に相補的なsiRNAと複合体を形成してその領域のメチル化ヒストンと安定に結合し、RNAポリメラーゼによって転写が開始されたpre-mRNA新生鎖を、転写と共役した形で分解している。ヘテロクロマチン領域の維持ではなく形成過程はDicer依存的であるが、それはおそらく転写産物を標的とするsiRNAが最初に形成される際にはDicerが必要であるためである<a href="#cite_note-Noma-56">[56]</a>。新たなsiRNAは偶発的な転写による新生鎖からRdRPによって形成され、その領域に位置するRITS複合体へ取り込まれるため、ヘテロクロマチンの維持は自己強化型フィードバックループとして機能することが示唆されている<a href="#cite_note-Sugiyama-57">[57]</a>。分裂酵母の接合型遺伝子座やセントロメアにおける観察と哺乳類での現象との対応は明らかではなく、哺乳類細胞におけるヘテロクロマチンの維持はRNAi経路の構成要素とは無関係である可能性もある<a href="#cite_note-Wang_2006-58">[58]</a>。 <div class="mw-heading mw-heading3"><h3 id="RNA編集とのクロストーク">RNA編集とのクロストーク</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=6" title="節を編集: RNA編集とのクロストーク">編集</a>]</div> 高等真核生物で最も広くみられる<a href="/wiki/RNA%E7%B7%A8%E9%9B%86" class="mw-redirect" title="RNA編集">RNA編集</a>は、<a href="/wiki/ADAR" title="ADAR">ADAR</a>によるdsRNA中の<a href="/wiki/%E3%82%A2%E3%83%87%E3%83%8E%E3%82%B7%E3%83%B3" title="アデノシン">アデノシン</a>ヌクレオチド（A）の<a href="/wiki/%E3%82%A4%E3%83%8E%E3%82%B7%E3%83%B3" title="イノシン">イノシン</a>（I）への変換である<a href="#cite_note-Bass_2002-59">[59]</a>。RNAiとA→IのRNA編集経路が共通したdsRNA基質をめぐって競合する可能性は2000年に提唱された<a href="#cite_note-Bass_2000-60">[60]</a>。一部のpre-miRNAはA→I RNA編集を受け<a href="#cite_note-Luciano_2004-61">[61]</a><a href="#cite_note-yang2006-62">[62]</a>、この機構は成熟型miRNAへのプロセシングと発現を調節している可能性がある<a href="#cite_note-yang2006-62">[62]</a>。さらに、哺乳類のADARの少なくとも1種類に関してはsiRNAをRNAi経路の構成要素から隔離することが示されている<a href="#cite_note-Yang_2005-63">[63]</a>。ADARを持たないC. elegans系統を用いた研究からは、内在性遺伝子や導入遺伝子のRNAiによるサイレンシングにA→I RNA編集が対抗していることが示されており、このこともこのモデルを支持している<a href="#cite_note-Nishikura_2006-64">[64]</a>。<figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Rnai_diagram_retrovirology.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_diagram_retrovirology.png/300px-Rnai_diagram_retrovirology.png" decoding="async" width="300" height="295" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_diagram_retrovirology.png/450px-Rnai_diagram_retrovirology.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_diagram_retrovirology.png/600px-Rnai_diagram_retrovirology.png 2x" data-file-width="1200" data-file-height="1179" /></a><figcaption>植物と動物における遺伝子サイレンシグの大きな差異。内因性に発現したmiRNAまたは外因性のsiRNAはDicerによってプロセシングされてRISCに取り込まれ、遺伝子サイレンシングを媒介する<a href="#cite_note-Saumet-65">[65]</a>。</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="生物種間の多様性">生物種間の多様性</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=7" title="節を編集: 生物種間の多様性">編集</a>]</div> 外来dsRNAを取り込み、そしてそれらをRNAi経路で利用する能力は、生物によって差がある。RNAiの効果は、植物やC. eleganでは全身的かつ遺伝性のものであるが、ショウジョウバエや哺乳類ではそうではない。植物では、RNAiは<a href="/wiki/%E5%8E%9F%E5%BD%A2%E8%B3%AA%E9%80%A3%E7%B5%A1" title="原形質連絡">原形質連絡</a>（細胞間のコミュニケーションや輸送を可能にする、<a href="/wiki/%E7%B4%B0%E8%83%9E%E5%A3%81" title="細胞壁">細胞壁</a>に存在するチャネル）を介した細胞間でのsiRNAの輸送によって全身へ伝播していくと考えられている<a href="#cite_note-Lodish-36">[36]</a>。またその遺伝性はRNAiの標的となったプロモーターがメチル化されることによるものであり、メチル化パターンは新たな世代の細胞が生じるたびにコピーされる<a href="#cite_note-:10-66">[66]</a>。植物と動物のおおまかな違いは、内因的に産生されるmiRNAの標的性にある。植物では、通常miRNAはその標的遺伝子に対して（ほぼ）完全に相補的であり、RISCによる直接的なmRNAの切断が誘導されるのに対し、動物のmiRNAは標的となる配列がより多様である傾向があり、翻訳抑制が誘導される<a href="#cite_note-Saumet-65">[65]</a>。この翻訳抑制効果は、<a href="/wiki/%E7%9C%9F%E6%A0%B8%E7%94%9F%E7%89%A9%E3%81%AE%E7%BF%BB%E8%A8%B3%E9%96%8B%E5%A7%8B%E5%9B%A0%E5%AD%90" title="真核生物の翻訳開始因子">翻訳開始因子</a>とmRNAの<a href="/wiki/%E3%83%9D%E3%83%AA%E3%82%A2%E3%83%87%E3%83%8B%E3%83%AB%E5%8C%96" title="ポリアデニル化">ポリアデニル化</a>テールとの相互作用の阻害によって行われている可能性がある<a href="#cite_note-Humphreys-67">[67]</a>。 <a href="/wiki/%E3%83%AA%E3%83%BC%E3%82%B7%E3%83%A5%E3%83%9E%E3%83%8B%E3%82%A2" title="リーシュマニア">リーシュマニア</a>Leishmania majorや<a href="/wiki/%E3%83%88%E3%83%AA%E3%83%91%E3%83%8E%E3%82%BD%E3%83%BC%E3%83%9E" title="トリパノソーマ">トリパノソーマ</a><a href="/wiki/Trypanosoma_cruzi" class="mw-redirect" title="Trypanosoma cruzi">Trypanosoma cruzi</a>など、一部の原生動物にはRNAi経路が全く存在しない<a href="#cite_note-DaRocha_2004-68">[68]</a><a href="#cite_note-Robinson_2003-69">[69]</a>。一部の<a href="/wiki/%E8%8F%8C%E9%A1%9E" title="菌類">菌類</a>でも大部分またはすべての構成要素が存在せず、そうした生物として最も有名なのは<a href="/wiki/%E3%83%A2%E3%83%87%E3%83%AB%E7%94%9F%E7%89%A9" title="モデル生物">モデル生物</a>でもある<a href="/wiki/%E5%87%BA%E8%8A%BD%E9%85%B5%E6%AF%8D" title="出芽酵母">出芽酵母</a>Saccharomyces cerevisiaeである<a href="#cite_note-Aravind-70">[70]</a>。Saccharomyces castelliiや<a href="/wiki/%E3%82%AB%E3%83%B3%E3%82%B8%E3%83%80" title="カンジダ">カンジダ</a>Candida albicansなどの他の出芽酵母にはRNAiが存在し、S. castellii由来の2つのRNAi関連タンパク質を誘導することで、S. cerevisiaeでもRNAiが促進されることが示されている<a href="#cite_note-:11-71">[71]</a>。<a href="/wiki/%E5%AD%90%E5%9A%A2%E8%8F%8C%E9%96%80" title="子嚢菌門">子嚢菌門</a>や<a href="/wiki/%E6%8B%85%E5%AD%90%E8%8F%8C%E9%96%80" title="担子菌門">担子菌門</a>の特定の種でRNAi経路が存在しないことは、RNAサイレンシングに必要なタンパク質が多くの菌類系統で独立に失われたことを示しており、おそらく類似した機能を持つ新たな経路の進化か、特定の<a href="/wiki/%E3%83%8B%E3%83%83%E3%83%81" title="ニッチ">ニッチ</a>において選択上の有利さが存在しなかったためであると考えられる<a href="#cite_note-Nakayashiki-72">[72]</a>。 <div class="mw-heading mw-heading3"><h3 id="関連する原核生物のシステム">関連する原核生物のシステム</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=8" title="節を編集: 関連する原核生物のシステム">編集</a>]</div> <a href="/wiki/%E5%8E%9F%E6%A0%B8%E7%94%9F%E7%89%A9" title="原核生物">原核生物</a>における遺伝子発現は、いくつかの面でRNAiと類似したRNAベースの系の影響を受ける。RNAをコードする遺伝子は、mRNAとアニーリングする相補的なRNAを産生することで、mRNAの存在量や翻訳を制御する。しかし、こうした調節性RNAにはDicerは関与せず、一般的にはmiRNAに類似したものとはみなされていない<a href="#cite_note-Morita_2006-73">[73]</a>。原核生物における<a href="/wiki/CRISPR" title="CRISPR">CRISPR</a>系が真核生物におけるRNAi系と類似したものであることも示唆されているが、どのタンパク質構成要素も<a href="/wiki/%E3%82%AA%E3%83%BC%E3%82%BD%E3%83%AD%E3%82%AC%E3%82%B9" class="mw-redirect" title="オーソロガス">オーソロガス</a>ではない<a href="#cite_note-makarova-74">[74]</a>。 <div class="mw-heading mw-heading2"><h2 id="生物学的機能">生物学的機能</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=9" title="節を編集: 生物学的機能">編集</a>]</div> <div class="mw-heading mw-heading3"><h3 id="免疫">免疫</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=10" title="節を編集: 免疫">編集</a>]</div> RNAiはウイルスやその他の外来性遺伝物質に対する<a href="/wiki/%E5%85%8D%E7%96%AB%E5%BF%9C%E7%AD%94" title="免疫応答">免疫応答</a>の重要な部分をなしており、特に植物では<a href="/wiki/%E3%83%88%E3%83%A9%E3%83%B3%E3%82%B9%E3%83%9D%E3%82%BE%E3%83%B3" title="トランスポゾン">トランスポゾン</a>の自己増殖も防いでいる<a href="#cite_note-Stram_2006-75">[75]</a>。<a href="/wiki/%E3%82%B7%E3%83%AD%E3%82%A4%E3%83%8C%E3%83%8A%E3%82%BA%E3%83%8A" title="シロイヌナズナ">シロイヌナズナ</a>Arabidopsis thalianaなどの植物は複数のDicerホモログを発現しており、これらは植物が異なるウイルスに曝露した際には異なる反応を示すよう専門化されている<a href="#cite_note-Blevins-76">[76]</a>。RNAi経路が十分に理解される前から、植物における遺伝子サイレンシングの誘導がその植物全体に伝播したり、また<a href="/wiki/%E6%8E%A5%E3%81%8E%E6%9C%A8" title="接ぎ木">接ぎ木</a>によって台木から接ぎ穂へ移行したりすることがあることが知られていた<a href="#cite_note-Palauqui-77">[77]</a>。この現象は、ウイルスに最初に遭遇した後、植物全体がウイルスへ応答することを可能にする、植物の免疫系の特徴として認識されていた<a href="#cite_note-Voinnet-78">[78]</a>。一方、多くの植物ウイルスもRNAi経路を抑制する精巧な機構を進化させてきた<a href="#cite_note-Lucy-79">[79]</a>。こうしたものの中には、Dicerによって産生される一本鎖オーバーハング末端を持つ短い二本鎖RNA断片に対して結合するウイルスタンパク質などが含まれる<a href="#cite_note-Merai-80">[80]</a>。一部の植物のゲノムは、特定種の<a href="/wiki/%E7%B4%B0%E8%8F%8C" title="細菌">細菌</a>の感染に対する応答として、内因性のsiRNAを発現する<a href="#cite_note-Katiyar-Agarwal-81">[81]</a>。これらは、感染過程の助けとなりうる宿主のあらゆる代謝過程をダウンレギュレーションする、病原体に対する全般的応答の一部である可能性がある<a href="#cite_note-Fritz-82">[82]</a>。 一般的に動物で発現しているDicerの種類は植物よりも少ないが、一部の動物ではRNAiが抗ウイルス応答を行っている。ショウジョウバエでは幼体と成体の双方において、RNAiは抗ウイルス免疫応答に重要であり、<a href="/wiki/%E3%82%B7%E3%83%A7%E3%82%A6%E3%82%B8%E3%83%A7%E3%82%A6%E3%83%90%E3%82%A8X%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ショウジョウバエXウイルス">ショウジョウバエXウイルス</a>などの病原体に対して活性を示す<a href="#cite_note-Zambon-83">[83]</a><a href="#cite_note-Wang-84">[84]</a>。免疫における同様の役割はC. elegansでも作用している可能性があり、ウイルスに応答してArgonauteタンパク質はアップレギュレーションされ、またRNAi経路のの構成要素を過剰発現した線虫はウイルス感染に対して耐性を示す<a href="#cite_note-Lu-85">[85]</a><a href="#cite_note-Wilkins-86">[86]</a>。 哺乳類の<a href="/wiki/%E8%87%AA%E7%84%B6%E5%85%8D%E7%96%AB%E7%B3%BB" title="自然免疫系">自然免疫</a>におけるRNAiの役割の理解は進んでおらず、比較的わずかなデータしか存在しない。哺乳類細胞における機能的な抗ウイルスRNAi経路の存在を示す証拠は提示されており<a href="#cite_note-:12-87">[87]</a><a href="#cite_note-:13-88">[88]</a>、また哺乳類細胞のRNAi応答を抑制する遺伝子をコードするウイルスの存在は哺乳類におけるRNAi依存的な免疫応答を支持する証拠となる可能性がある<a href="#cite_note-Berkhout-89">[89]</a><a href="#cite_note-Schutz-90">[90]</a>。しかしながら、この仮説には十分な証拠がないとして異議も唱えられている<a href="#cite_note-Cullen-91">[91]</a>。 哺乳類のウイルスにおけるRNAiには他の機能もあり、<a href="/wiki/%E3%83%98%E3%83%AB%E3%83%9A%E3%82%B9%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ヘルペスウイルス">ヘルペスウイルス</a>が発現するmiRNAはウイルス潜伏を媒介するヘテロクロマチン組織化の引き金として作用している可能性がある<a href="#cite_note-Li3-92">[92]</a>。 <div class="mw-heading mw-heading3"><h3 id="遺伝子のダウンレギュレーション">遺伝子のダウンレギュレーション</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=11" title="節を編集: 遺伝子のダウンレギュレーション">編集</a>]</div> 内因的に発現しているmiRNAは、<a href="/wiki/%E3%82%A4%E3%83%B3%E3%83%88%E3%83%AD%E3%83%B3" title="イントロン">イントロン</a>内に位置するのものも遺伝子間領域に位置するものも、翻訳抑制<a href="#cite_note-Saumet-65">[65]</a>そして発生の調節に最も重要であり、<a href="/wiki/%E5%BD%A2%E6%85%8B%E5%BD%A2%E6%88%90" title="形態形成">形態形成</a>の時期の決定や、<a href="/wiki/%E5%B9%B9%E7%B4%B0%E8%83%9E" title="幹細胞">幹細胞</a>などの未分化状態や不完全分化状態の細胞種の維持に特に重要である<a href="#cite_note-Carrington-93">[93]</a>。遺伝子発現のダウンレギュレーションにおける内因性miRNAの役割は、1993年にC. elegansで初めて記載された<a href="#cite_note-Lee_1993-94">[94]</a>。植物では、この機能はシロイヌナズナのmiR-JAWが植物の形状を制御するいくつかの遺伝子の調節に関与していることが示された際に発見された<a href="#cite_note-Palatnik_2003-95">[95]</a>。植物では、miRNAによって調節される遺伝子の大部分は<a href="/wiki/%E8%BB%A2%E5%86%99%E5%9B%A0%E5%AD%90" title="転写因子">転写因子</a>である<a href="#cite_note-Zhang_plant-96">[96]</a>。そのためmiRNAの活性は特に広範囲にわたり、転写因子や<a href="/wiki/F%E3%83%9C%E3%83%83%E3%82%AF%E3%82%B9%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA" title="Fボックスタンパク質">Fボックスタンパク質</a>など重要な調節遺伝子を調節することで発生時に遺伝子ネットワーク全体を調節する<a href="#cite_note-Jones-Rhoades-97">[97]</a>。ヒトを含む多くの生物では、miRNAは<a href="/wiki/%E8%85%AB%E7%98%8D" title="腫瘍">腫瘍</a>形成や<a href="/wiki/%E7%B4%B0%E8%83%9E%E5%91%A8%E6%9C%9F" title="細胞周期">細胞周期</a>の調節異常と関連づけられている。miRNAは<a href="/wiki/%E3%81%8C%E3%82%93%E9%81%BA%E4%BC%9D%E5%AD%90" title="がん遺伝子">がん遺伝子</a>としても<a href="/wiki/%E3%81%8C%E3%82%93%E6%8A%91%E5%88%B6%E9%81%BA%E4%BC%9D%E5%AD%90" title="がん抑制遺伝子">がん抑制遺伝子</a>としても機能する<a href="#cite_note-Zhang_cancer-98">[98]</a>。 <div class="mw-heading mw-heading2"><h2 id="進化">進化</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=12" title="節を編集: 進化">編集</a>]</div> <a href="/wiki/%E6%9C%80%E5%A4%A7%E7%AF%80%E7%B4%84%E6%B3%95" title="最大節約法">最大節約法</a>による<a href="/wiki/%E7%B3%BB%E7%B5%B1%E5%AD%A6" title="系統学">系統学</a>的解析に基づくと、全真核生物の<a href="/wiki/%E6%9C%80%E3%82%82%E8%BF%91%E3%81%84%E5%85%B1%E9%80%9A%E7%A5%96%E5%85%88" title="最も近い共通祖先">最近共通祖先</a>はすでに原始的なRNAi経路を持っていた可能性が極めて高く、特定の真核生物にRNAi経路が存在しないことは派生形質であると考えられている<a href="#cite_note-Cerutti-99">[99]</a>。この祖先型のRNAi系は、少なくとも1つのDicer様タンパク質、1つのArgonauteタンパク質、1つの<a href="/w/index.php?title=Piwi&action=edit&redlink=1" class="new" title="「Piwi」 (存在しないページ)">Piwi</a>（<a href="https://en.wikipedia.org/wiki/Piwi" class="extiw" title="en:Piwi">英語版</a>）タンパク質、そしてRNA依存性RNAポリメラーゼを含んでいたと考えられ、これらは細胞中で他の役割を果たしていた可能性もある。大規模な<a href="/wiki/%E6%AF%94%E8%BC%83%E3%82%B2%E3%83%8E%E3%83%9F%E3%82%AF%E3%82%B9" title="比較ゲノミクス">比較ゲノミクス</a>研究でも同様に、真核生物の<a href="/wiki/%E3%82%AF%E3%83%A9%E3%82%A6%E3%83%B3%E3%82%B0%E3%83%AB%E3%83%BC%E3%83%97" title="クラウングループ">クラウングループ</a>はすでにこれらの構成要素を持っており、<a href="/wiki/%E3%82%A8%E3%82%AD%E3%82%BD%E3%82%BD%E3%83%BC%E3%83%A0%E8%A4%87%E5%90%88%E4%BD%93" title="エキソソーム複合体">エキソソーム</a>などの一般的なRNA分解系とより密接な機能的関係を持っていた可能性があることが示唆されている<a href="#cite_note-Anantharaman-100">[100]</a>。またこの研究は、真核生物、大部分の古細菌、そして少なくとも一部の細菌（<a href="/w/index.php?title=Aquifex_aeolicus&action=edit&redlink=1" class="new" title="「Aquifex aeolicus」 (存在しないページ)">Aquifex aeolicus</a>（<a href="https://en.wikipedia.org/wiki/Aquifex_aeolicus" class="extiw" title="en:Aquifex aeolicus">英語版</a>）など）に共通して存在するRNA結合性のArgonauteタンパク質ファミリーは、翻訳開始系の構成要素と相同であり、そしてそこから進化したものであることも示唆している<a href="#cite_note-Anantharaman-100">[100]</a>。 <div class="mw-heading mw-heading2"><h2 id="応用">応用</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=13" title="節を編集: 応用">編集</a>]</div> <div class="mw-heading mw-heading3"><h3 id="遺伝子ノックダウンのためのRNAi経路">遺伝子ノックダウンのためのRNAi経路</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=14" title="節を編集: 遺伝子ノックダウンのためのRNAi経路">編集</a>]</div> RNAi経路は実験生物学において、培養細胞やモデル生物におけるin vivoでの遺伝子機能の研究のためにしばしば利用される<a href="#cite_note-Daneholt2006-34">[34]</a>。目的の遺伝子に対して相補的な配列を持つ二本鎖RNAが合成され、細胞または個体に導入される。そこで二本鎖RNAは外因性の遺伝子物質と認識され、RNAi経路が活性化される。この機構を用いて標的遺伝子の発現の劇的な低下を引き起こすことが可能であり、そしてこの低下の影響を研究することで遺伝子産物の生理学的役割を示すことができる。RNAiでは遺伝子の発現が完全には抑制されない場合があるため、遺伝子の発現が完全に除去される「<a href="/wiki/%E9%81%BA%E4%BC%9D%E5%AD%90%E3%83%8E%E3%83%83%E3%82%AF%E3%82%A2%E3%82%A6%E3%83%88" title="遺伝子ノックアウト">ノックアウト</a>」と区別して、「<a href="/wiki/%E9%81%BA%E4%BC%9D%E5%AD%90%E3%83%8E%E3%83%83%E3%82%AF%E3%83%80%E3%82%A6%E3%83%B3" title="遺伝子ノックダウン">ノックダウン</a>」と呼ばれることがある<a href="#cite_note-pmid12480342-101">[101]</a>。遺伝子アレイデータを用いたRNAiのサイレンシング効率の検証では、429の独立した実験において失敗率は18.5%であることが示されている<a href="#cite_note-:14-102">[102]</a>。 遺伝子ノックダウン効果を最大化しオフターゲット効果を最小化するdsRNAの設計法に関して、<a href="/wiki/%E8%A8%88%E7%AE%97%E7%94%9F%E7%89%A9%E5%AD%A6" title="計算生物学">計算生物学</a>では多くの研究がなされている。オフターゲット効果は、導入されたRNAの配列が複数の遺伝子と対合して発現を低下させるために生じる。ヒト、C. elegans、S. pombeのゲノム研究からは、可能なsiRNA配列のうち約10%で重大なオフターゲット効果が生じる可能性があると推定されている<a href="#cite_note-Qiu-7">[7]</a>。一般的<a href="#cite_note-Naito_ds-103">[103]</a><a href="#cite_note-Henschel-104">[104]</a>、哺乳類特異的<a href="#cite_note-Naito_si-105">[105]</a>、そしてウイルス特異的<a href="#cite_note-Naito_virus-106">[106]</a>なsiRNAを設計し、自動的に交差反応性のチェックを行うアルゴリズムを備えたソフトウェアツールが多く開発されている。 生物種や実験系に応じて、外因性RNAはDicerによって切断されるように設計された長鎖RNAである場合や、siRNA基質として作用するよう設計された短鎖RNAである場合がある。ほとんどの哺乳類細胞では、長い二本鎖RNA分子に対しては外来遺伝物質に非特異的に作用する<a href="/wiki/%E8%87%AA%E7%84%B6%E5%85%8D%E7%96%AB%E7%B3%BB" title="自然免疫系">自然免疫</a>の一種である<a href="/wiki/%E3%82%A4%E3%83%B3%E3%82%BF%E3%83%BC%E3%83%95%E3%82%A7%E3%83%AD%E3%83%B3" title="インターフェロン">インターフェロン</a>応答が誘導されるため、短いRNAが利用される<a href="#cite_note-Reynolds-107">[107]</a>。マウスの<a href="/wiki/%E5%8D%B5%E6%AF%8D%E7%B4%B0%E8%83%9E" title="卵母細胞">卵母細胞</a>や初期<a href="/wiki/%E8%83%9A" title="胚">胚</a>は外因性dsRNAに対するこの応答を欠くため、哺乳類の遺伝子ノックダウン効果を研究するための一般的なモデル系となっている<a href="#cite_note-Stein_oocyte-108">[108]</a>。siRNAが転写される適切な配列をコードしたプラスミドの安定トランスフェクションや<a href="#cite_note-Brummelkamp-109">[109]</a>、より精巧な<a href="/wiki/%E3%83%AC%E3%83%B3%E3%83%81%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9%E5%B1%9E" title="レンチウイルス属">レンチウイルス</a>ベクターシステムによって転写の活性化や不活性化の誘導を可能にしたコンディショナルRNAi（conditional RNAi）と呼ばれる技術など<a href="#cite_note-Tiscornia-110">[110]</a><a href="#cite_note-Ventura-111">[111]</a>、siRNAの直接導入を回避することで哺乳類系におけるRNAiの有用性を改善した特殊な実験技術も開発されている。 <div class="mw-heading mw-heading3"><h3 id="機能ゲノミクス">機能ゲノミクス</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=15" title="節を編集: 機能ゲノミクス">編集</a>]</div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Drosophila_melanogaster_-_side_(aka).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/250px-Drosophila_melanogaster_-_side_%28aka%29.jpg" decoding="async" width="250" height="194" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/375px-Drosophila_melanogaster_-_side_%28aka%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/500px-Drosophila_melanogaster_-_side_%28aka%29.jpg 2x" data-file-width="546" data-file-height="424" /></a><figcaption>RNAi実験で一般的に利用されるモデル生物であるショウジョウバエの正常な成体</figcaption></figure> ゲノムワイドRNAiライブラリの設計には、一定の実験条件のセットに対して単一のsiRNAを設計するよりも高度な技術が必要となる場合がある。siRNAライブラリの設計<a href="#cite_note-Huesken-112">[112]</a>や遺伝子ノックダウン時の効率の予測<a href="#cite_note-Ge-113">[113]</a>には、<a href="/wiki/%E3%83%8B%E3%83%A5%E3%83%BC%E3%83%A9%E3%83%AB%E3%83%8D%E3%83%83%E3%83%88%E3%83%AF%E3%83%BC%E3%82%AF" title="ニューラルネットワーク">ニューラルネットワーク</a>がよく利用される。マスゲノムスクリーニングは<a href="/w/index.php?title=DNA%E3%82%A2%E3%83%8E%E3%83%86%E3%83%BC%E3%82%B7%E3%83%A7%E3%83%B3&action=edit&redlink=1" class="new" title="「DNAアノテーション」 (存在しないページ)">ゲノムアノテーション</a>（<a href="https://en.wikipedia.org/wiki/DNA_annotation" class="extiw" title="en:DNA annotation">英語版</a>）のための有望な方法として広く知られており、<a href="/wiki/%E3%83%9E%E3%82%A4%E3%82%AF%E3%83%AD%E3%82%A2%E3%83%AC%E3%82%A4" title="マイクロアレイ">マイクロアレイ</a>ベースの<a href="/wiki/%E3%83%8F%E3%82%A4%E3%82%B9%E3%83%AB%E3%83%BC%E3%83%97%E3%83%83%E3%83%88%E3%82%B9%E3%82%AF%E3%83%AA%E3%83%BC%E3%83%8B%E3%83%B3%E3%82%B0" title="ハイスループットスクリーニング">ハイスループットスクリーニング</a>の開発の引き金となった<a href="#cite_note-Janitz-114">[114]</a><a href="#cite_note-Vanhecke-115">[115]</a>。しかし、こうしたスクリーニングの有用性や、モデル生物で開発された技術が近縁種にも一般化できるのか、例えばC. elegansの技術が関連する寄生性線虫に応用可能であるのかに関しては疑問視されている<a href="#cite_note-Geldhof-116">[116]</a><a href="#cite_note-Geldhof2-117">[117]</a>。 <div class="mw-heading mw-heading3"><h3 id="医療">医療</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=16" title="節を編集: 医療">編集</a>]</div> 医療におけるRNAiの利用の歴史 <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf/page1-330px-Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf.jpg" decoding="async" width="330" height="427" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf/page1-495px-Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/59/Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf/page1-660px-Vertical_Timeline_of_RNAi%27s_Use_in_Medicine.pdf.jpg 2x" data-file-width="1275" data-file-height="1650" /></a><figcaption>医療におけるRNAiの利用の時系列</figcaption></figure> 動物におけるRNAサイレンシングの最初の例は1996年に記載された。線虫C. elegansにおいて、par-1 mRNAのセンス鎖とアンチセンス鎖のRNAを導入することで、par-1 mRNAの分解が引き起こされることが観察された<a href="#cite_note-Sen-118">[118]</a>。この分解は一本鎖RNAによって開始されると考えられていたが、その2年後の1998年、ファイアーとメローによってこのpar-1遺伝子発現のサイレンシング能力は実際には二本鎖RNAによって開始されていることが発見された<a href="#cite_note-Sen-118">[118]</a>。この発見によって、彼らはノーベル生理学・医学賞を受賞した<a href="#cite_note-:15-119">[119]</a>。この画期的な発見の直後、合成siRNAを用いることで、サイレンシングの標的は遺伝子全体ではなく遺伝子中の特定の配列とすることが可能であることが発見された<a href="#cite_note-:16-120">[120]</a>。そのわずか1年後、トランスジェニックマウスにおいて<a href="/wiki/C%E5%9E%8B%E8%82%9D%E7%82%8E%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="C型肝炎ウイルス">C型肝炎ウイルス</a>配列を標的として、この配列特異的なサイレンシングの治療応用が実証された<a href="#cite_note-:17-121">[121]</a>。それ以降、RNAiの治療応用を広げる試みが多くの研究者によってなされ、具体的にはさまざまなタイプのがんを引き起こす遺伝子を標的とすることに関心が寄せられた<a href="#cite_note-:18-122">[122]</a><a href="#cite_note-:19-123">[123]</a>。2006年までに臨床試験に到達した最初の応用は、<a href="/wiki/%E5%8A%A0%E9%BD%A2%E9%BB%84%E6%96%91%E5%A4%89%E6%80%A7" title="加齢黄斑変性">黄斑変性</a>と<a href="/wiki/RS%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="RSウイルス">RSウイルス</a>の治療であった<a href="#cite_note-:20-124">[124]</a>。その4年後、ナノ粒子デリバリーシステムを用いて、固形腫瘍を標的としたヒトでの初めての第I相臨床試験が開始された<a href="#cite_note-:21-125">[125]</a>。現在のところ大部分の研究はがん治療へのRNAiの応用を試みるものであるが、可能な応用は広範囲にわたる。RNAiは、<a href="/wiki/%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ウイルス">ウイルス</a><a href="#cite_note-:22-126">[126]</a>、<a href="/wiki/%E7%B4%B0%E8%8F%8C" title="細菌">細菌</a><a href="#cite_note-:23-127">[127]</a>、<a href="/wiki/%E5%AF%84%E7%94%9F%E8%99%AB" title="寄生虫">寄生虫</a>による感染症<a href="#cite_note-:24-128">[128]</a>、不適応な遺伝子変異の治療<a href="#cite_note-:25-129">[129]</a>、<a href="/wiki/%E8%96%AC%E7%89%A9%E4%B9%B1%E7%94%A8" title="薬物乱用">薬物使用</a>の制御<a href="#cite_note-:26-130">[130]</a>、<a href="/wiki/%E7%96%BC%E7%97%9B" class="mw-redirect" title="疼痛">疼痛</a>管理<a href="#cite_note-:27-131">[131]</a>、さらには睡眠の調節<a href="#cite_note-:28-132">[132]</a>にも利用できる可能性がある。 <div class="mw-heading mw-heading4"><h4 id="治療応用">治療応用</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=17" title="節を編集: 治療応用">編集</a>]</div> <div class="mw-heading mw-heading5"><h5 id="ウイルス感染">ウイルス感染</h5>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=18" title="節を編集: ウイルス感染">編集</a>]</div> 抗ウイルス治療はRNAiベースの医療応用として最初期に提唱されたものであり、2つの異なる種類のものが開発されている。1つはウイルスのRNAを標的とするものである。ウイルスのRNAを標的化することで、<a href="/wiki/%E3%83%92%E3%83%88%E5%85%8D%E7%96%AB%E4%B8%8D%E5%85%A8%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ヒト免疫不全ウイルス">HIV</a><a href="#cite_note-:29-133">[133]</a>、<a href="/wiki/%E3%83%92%E3%83%88%E3%83%91%E3%83%94%E3%83%AD%E3%83%BC%E3%83%9E%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="ヒトパピローマウイルス">HPV</a><a href="#cite_note-:30-134">[134]</a>、<a href="/wiki/A%E5%9E%8B%E8%82%9D%E7%82%8E%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="A型肝炎ウイルス">A型肝炎ウイルス</a><a href="#cite_note-:31-135">[135]</a>、<a href="/wiki/B%E5%9E%8B%E8%82%9D%E7%82%8E%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="B型肝炎ウイルス">B型肝炎ウイルス</a><a href="#cite_note-:32-136">[136]</a>、<a href="/wiki/%E3%82%A4%E3%83%B3%E3%83%95%E3%83%AB%E3%82%A8%E3%83%B3%E3%82%B6%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="インフルエンザウイルス">インフルエンザウイルス</a><a href="#cite_note-:33-137">[137]</a><a href="#cite_note-:34-138">[138]</a><a href="#cite_note-:35-139">[139]</a>、<a href="/wiki/RS%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="RSウイルス">RSウイルス</a><a href="#cite_note-:0-140">[140]</a>、<a href="/wiki/SARS%E3%82%B3%E3%83%AD%E3%83%8A%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="SARSコロナウイルス">SARSコロナウイルス</a><a href="#cite_note-:0-140">[140]</a>、<a href="/wiki/%E3%82%A2%E3%83%87%E3%83%8E%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="アデノウイルス">アデノウイルス</a><a href="#cite_note-:0-140">[140]</a>、<a href="/wiki/%E9%BA%BB%E7%96%B9%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="麻疹ウイルス">麻疹ウイルス</a><a href="#cite_note-:36-141">[141]</a>など、多数のウイルスの複製を抑制できることが多くの研究により示されている。もう1つの戦略は、宿主細胞の遺伝子を標的とすることでウイルスの侵入を防ぐものである<a href="#cite_note-:37-142">[142]</a>。たとえば、宿主細胞の<a href="/wiki/%E3%82%B1%E3%83%A2%E3%82%AB%E3%82%A4%E3%83%B3%E5%8F%97%E5%AE%B9%E4%BD%93" title="ケモカイン受容体">ケモカイン受容体</a>（<a href="/w/index.php?title=CXCR4&action=edit&redlink=1" class="new" title="「CXCR4」 (存在しないページ)">CXCR4</a>（<a href="https://en.wikipedia.org/wiki/CXCR4" class="extiw" title="en:CXCR4">英語版</a>）と<a href="/wiki/CCR5" title="CCR5">CCR5</a>）を抑制することでHIVの侵入を防ぐことができる<a href="#cite_note-:38-143">[143]</a>。 <div class="mw-heading mw-heading5"><h5 id="がん">がん</h5>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=19" title="節を編集: がん">編集</a>]</div> 伝統的な<a href="/wiki/%E5%8C%96%E5%AD%A6%E7%99%82%E6%B3%95_(%E6%82%AA%E6%80%A7%E8%85%AB%E7%98%8D)" title="化学療法 (悪性腫瘍)">化学療法</a>はがん細胞を効果的に殺すことができるが、正常細胞とがん細胞を区別する特異性を欠くために通常は重大な副作用を伴う。RNAiはがんと関連した遺伝子（<a href="/wiki/%E3%81%8C%E3%82%93%E9%81%BA%E4%BC%9D%E5%AD%90" title="がん遺伝子">がん遺伝子</a>など）を標的とすることで腫瘍の成長を阻害する、より特異的なアプローチとなることが多くの研究で示されている<a href="#cite_note-:39-144">[144]</a>。RNAiはがん細胞の化学療法薬に対する感受性を高めることも可能であり、化学療法との併用療法も提唱されている<a href="#cite_note-:40-145">[145]</a>。細胞の浸潤や<a href="/wiki/%E9%81%8A%E8%B5%B0" title="遊走">遊走</a>の阻害も、RNAiベースの他の治療法となる可能性がある<a href="#cite_note-:41-146">[146]</a>。 <div class="mw-heading mw-heading5"><h5 id="神経疾患">神経疾患</h5>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=20" title="節を編集: 神経疾患">編集</a>]</div> RNAiは<a href="/wiki/%E7%A5%9E%E7%B5%8C%E5%A4%89%E6%80%A7%E7%96%BE%E6%82%A3" title="神経変性疾患">神経変性疾患</a>の治療法となる可能性が示されている。細胞やマウスでの研究では、<a href="/wiki/%E3%82%A2%E3%83%9F%E3%83%AD%E3%82%A4%E3%83%89%CE%B2" title="アミロイドβ">アミロイドβ</a>（Aβ）を産生する遺伝子（<a href="/wiki/%CE%92-%E3%82%BB%E3%82%AF%E3%83%AC%E3%82%BF%E3%83%BC%E3%82%BC1" title="Β-セクレターゼ1">BACE1</a>や<a href="/wiki/%E3%82%A2%E3%83%9F%E3%83%AD%E3%82%A4%E3%83%89%E5%89%8D%E9%A7%86%E4%BD%93%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA" title="アミロイド前駆体タンパク質">APP</a>など）をRNAiで特異的に標的化することで、<a href="/wiki/%E3%82%A2%E3%83%AB%E3%83%84%E3%83%8F%E3%82%A4%E3%83%9E%E3%83%BC%E7%97%85" title="アルツハイマー病">アルツハイマー病</a>と関係するAβペプチドの量を大きく低下させることができることが示されている<a href="#cite_note-:42-147">[147]</a><a href="#cite_note-:43-148">[148]</a><a href="#cite_note-:44-149">[149]</a>。さらに、こうしたサイレンシングベースのアプローチは、<a href="/wiki/%E3%83%91%E3%83%BC%E3%82%AD%E3%83%B3%E3%82%BD%E3%83%B3%E7%97%85" title="パーキンソン病">パーキンソン病</a>や<a href="/wiki/%E3%83%88%E3%83%AA%E3%83%97%E3%83%AC%E3%83%83%E3%83%88%E3%83%AA%E3%83%94%E3%83%BC%E3%83%88%E7%97%85" title="トリプレットリピート病">ポリグルタミン病</a>の治療においても有望な結果をもたらしている<a href="#cite_note-:45-150">[150]</a><a href="#cite_note-:46-151">[151]</a><a href="#cite_note-:47-152">[152]</a>。 <div class="mw-heading mw-heading4"><h4 id="治療応用における困難">治療応用における困難</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=21" title="節を編集: 治療応用における困難">編集</a>]</div> RNAiの臨床的可能性の実現のためには、siRNAが効率的に標的組織の細胞へ輸送される必要がある。しかしながら、臨床利用までに克服すべきさまざまな障壁が存在する。例えば、「裸」のsiRNAはその治療効力を低下させるいくつかの障害の影響を受けやすい<a href="#cite_note-Kanasty_nmat3765-153">[153]</a>。いったんsiRNAが血流に移行すると、裸のRNAは血清中の<a href="/wiki/%E3%83%8C%E3%82%AF%E3%83%AC%E3%82%A2%E3%83%BC%E3%82%BC" title="ヌクレアーゼ">ヌクレアーゼ</a>によって分解されたり、自然免疫系を刺激したりする<a href="#cite_note-Kanasty_nmat3765-153">[153]</a>。また、そのサイズと高い<a href="/wiki/%E3%82%A2%E3%83%8B%E3%82%AA%E3%83%B3" class="mw-redirect" title="アニオン">アニオン</a>性のため、未修飾のsiRNA分子が<a href="/wiki/%E7%B4%B0%E8%83%9E%E8%86%9C" title="細胞膜">細胞膜</a>を通って細胞内へ移行するのは容易ではない。そのため、人工的なsiRNAやナノ粒子に封入したsiRNAを利用する必要がある。しかしながら、細胞膜を越えたsiRNAの輸送にはさらに固有の問題が存在する。siRNAが細胞膜を越えて輸送された場合、その量が最適化されていなければ意図しない毒性が生じたり、オフターゲット効果（部分的な配列相補性を持つ遺伝子に対する意図しないダウンレギュレーションなど）が生じたりする可能性がある<a href="#cite_note-ReferenceA-154">[154]</a>。細胞に移行した後も、その効果は細胞分裂ごとに希釈されるため繰り返し投与が必要である。また、dsRNAを運搬するベクターの一部には調節作用がある場合があるため、非特異的な副作用も考慮し、制御される必要がある<a href="#cite_note-:48-155">[155]</a>。 <div class="mw-heading mw-heading4"><h4 id="免疫応答の刺激">免疫応答の刺激</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=22" title="節を編集: 免疫応答の刺激">編集</a>]</div> ヒトの免疫系は、<a href="/wiki/%E8%87%AA%E7%84%B6%E5%85%8D%E7%96%AB%E7%B3%BB" title="自然免疫系">自然免疫系</a>と<a href="/wiki/%E7%8D%B2%E5%BE%97%E5%85%8D%E7%96%AB%E7%B3%BB" title="獲得免疫系">獲得免疫系</a>の2種類に分類される<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。自然免疫系は感染に対する第一の防御機構であり、病原体に対して一般的応答を行う<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。一方、獲得免疫系は自然免疫系よりも後で進化した系であり、病原体の分子の特定の部分に反応するよう訓練された、高度に専門化された<a href="/wiki/B%E7%B4%B0%E8%83%9E" title="B細胞">B細胞</a>と<a href="/wiki/T%E7%B4%B0%E8%83%9E" title="T細胞">T細胞</a>によって構成される<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。 siRNAは自然免疫系によって制御されており、自然免疫系による応答はさらに急性炎症応答と抗ウイルス応答に分類される<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。炎症応答では、低分子シグナル伝達分子や<a href="/wiki/%E3%82%A4%E3%83%B3%E3%82%BF%E3%83%BC%E3%83%AD%E3%82%A4%E3%82%AD%E3%83%B3-1" title="インターロイキン-1">IL-1</a>、<a href="/wiki/%E3%82%A4%E3%83%B3%E3%82%BF%E3%83%BC%E3%83%AD%E3%82%A4%E3%82%AD%E3%83%B3-6" title="インターロイキン-6">IL-6</a>、<a href="/wiki/%E3%82%A4%E3%83%B3%E3%82%BF%E3%83%BC%E3%83%AD%E3%82%A4%E3%82%AD%E3%83%B3-12" title="インターロイキン-12">IL-12</a>、<a href="/wiki/TNF-%CE%B1" class="mw-redirect" title="TNF-α">TNF-α</a>などの<a href="/wiki/%E3%82%B5%E3%82%A4%E3%83%88%E3%82%AB%E3%82%A4%E3%83%B3" title="サイトカイン">サイトカイン</a>が誘導される。こうした炎症性サイトカインは<a href="/wiki/%E9%A3%9F%E4%BD%9C%E7%94%A8" title="食作用">食作用</a>を刺激し、侵入した病原体を破壊する<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。抗ウイルス応答では、IFN-αやIFN-βなどのタイプI<a href="/wiki/%E3%82%A4%E3%83%B3%E3%82%BF%E3%83%BC%E3%83%95%E3%82%A7%E3%83%AD%E3%83%B3" title="インターフェロン">インターフェロン</a>の放出や抗ウイルス遺伝子のアップレギュレーションが誘導される<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。どちらの応答も、<a href="/wiki/%E3%83%91%E3%82%BF%E3%83%BC%E3%83%B3%E8%AA%8D%E8%AD%98%E5%8F%97%E5%AE%B9%E4%BD%93" title="パターン認識受容体">パターン認識受容体</a>（PRR）の刺激を介して引き起こされる。複数のPRRによってRNA構造のさまざまな側面が認識されるため、免疫刺激を避けることは困難なものとなっている<a href="#cite_note-Whitehead_2011_77–96-156">[156]</a>。 <div class="mw-heading mw-heading4"><h4 id="治療技術としての展望">治療技術としての展望</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=23" title="節を編集: 治療技術としての展望">編集</a>]</div> 2015年から2017年にかけて行われたsiRNA治療の第I・II相試験では、肝臓での強力かつ持続的な遺伝子ノックダウン効果と臨床効果を示す一部の徴候がみられ、許容できない毒性はみられなかった<a href="#cite_note-ReferenceA-154">[154]</a>。<a href="/wiki/%E3%83%88%E3%83%A9%E3%83%B3%E3%82%B9%E3%82%B5%E3%82%A4%E3%83%AC%E3%83%81%E3%83%B3" title="トランスサイレチン">トランスサイレチン</a>の変異によって引き起こされる家族性神経変性・心筋症の治療へ向けた2つの第III相試験が進行中である<a href="#cite_note-ReferenceA-154">[154]</a>。多くの研究でin vivoデリバリーシステムの有望性は示されており、またそれらの多様な特性は無数の応用を可能にしている。最も有望なものはナノ粒子デリバリーシステムであるが、製品の安定した品質を確保するためには厳密に制御された混合過程が必要となることなど、製造過程のスケールアップにはさらなる課題が残されている<a href="#cite_note-Kanasty_nmat3765-153">[153]</a>。 <div class="mw-heading mw-heading3"><h3 id="バイオテクノロジー">バイオテクノロジー</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=24" title="節を編集: バイオテクノロジー">編集</a>]</div> RNAiは<a href="/wiki/%E3%83%90%E3%82%A4%E3%82%AA%E3%83%86%E3%82%AF%E3%83%8E%E3%83%AD%E3%82%B8%E3%83%BC" class="mw-redirect" title="バイオテクノロジー">バイオテクノロジー</a>分野で応用されており、他の分野でも商業化が近い。<a href="/wiki/%E3%83%8B%E3%82%B3%E3%83%81%E3%83%B3" title="ニコチン">ニコチン</a>を含まないタバコ、<a href="/wiki/%E3%82%AB%E3%83%95%E3%82%A7%E3%82%A4%E3%83%B3" title="カフェイン">カフェイン</a>を含まないコーヒー、栄養素を強化した野菜、低<a href="/wiki/%E3%82%A2%E3%83%AC%E3%83%AB%E3%82%B2%E3%83%B3" title="アレルゲン">アレルゲン</a>の作物など、RNAiを利用した新たな作物が発されている。遺伝子改変されたリンゴ<a href="/w/index.php?title=Arctic_Apples&action=edit&redlink=1" class="new" title="「Arctic Apples」 (存在しないページ)">Arctic Apples</a>（<a href="https://en.wikipedia.org/wiki/Arctic_Apples" class="extiw" title="en:Arctic Apples">英語版</a>）は2015年にFDAの承認を受けた<a href="#cite_note-:49-157">[157]</a>。このリンゴはPPO（<a href="/w/index.php?title=%E3%83%9D%E3%83%AA%E3%83%95%E3%82%A7%E3%83%8E%E3%83%BC%E3%83%AB%E3%82%AA%E3%82%AD%E3%82%B7%E3%83%80%E3%83%BC%E3%82%BC&action=edit&redlink=1" class="new" title="「ポリフェノールオキシダーゼ」 (存在しないページ)">ポリフェノールオキシダーゼ</a>（<a href="https://en.wikipedia.org/wiki/Polyphenol_oxidase" class="extiw" title="en:Polyphenol oxidase">英語版</a>））遺伝子をRNAiによって抑制することで、果実を切った後の褐変が起こらないようになっている。PPOがサイレンシングされたリンゴは、<a href="/wiki/%E3%82%AF%E3%83%AD%E3%83%AD%E3%82%B2%E3%83%B3%E9%85%B8" title="クロロゲン酸">クロロゲン酸</a>を標準的な<a href="/wiki/%E3%82%AD%E3%83%8E%E3%83%B3" title="キノン">キノン</a>産物へと変換することができないため、変色が起こらない<a href="#cite_note-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564-1">[1]</a>。 <a href="/wiki/%E4%BD%9C%E7%89%A9%E5%AD%A6" class="mw-redirect" title="作物学">作物学</a>におけるRNAiの応用には、ストレス耐性の付与や栄養素の強化などの改善などいくつかの可能性がある。RNAiはC3植物の生産性の向上のための光呼吸の阻害のほか、早期の開花、成熟や老化の遅れ、休眠の解除、ストレスに強い植物、自家不和合性の克服などの誘導に有用である可能性がある<a href="#cite_note-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564-1">[1]</a>。 <div class="mw-heading mw-heading4"><h4 id="食品">食品</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=25" title="節を編集: 食品">編集</a>]</div> RNAiは食料生産における将来的有望性が示されているが、まだ若い技術であるためその利点と欠点に対する理解に欠けるところがある。そのため、よりよく理解し誤解を取り除く必要がある<a href="#cite_note-Herrero-et-al-2020-158">[158]</a>。RNAiはすでに、天然毒素の産生が少ない遺伝子組み換え植物に利用されている。こうした技術は、植物においてRNAiの表現型が安定かつ遺伝性のものであることを利用している。<a href="/wiki/%E3%83%AF%E3%82%BF%E5%B1%9E" title="ワタ属">ワタ</a>の種子はタンパク質に富むが、有毒<a href="/wiki/%E3%83%86%E3%83%AB%E3%83%9A%E3%83%8E%E3%82%A4%E3%83%89" title="テルペノイド">テルペノイド</a>である<a href="/wiki/%E3%82%B4%E3%82%B7%E3%83%9D%E3%83%BC%E3%83%AB" title="ゴシポール">ゴシポール</a>を含むためヒトの食物としては適さない。ゴシポール自体は害虫による損傷から植物を守るために重要であるため、植物の他の部分に影響を与えることなく種子でのみゴシポールの産生に重要な酵素<a href="/w/index.php?title=%CE%94-%E3%82%AB%E3%82%B8%E3%83%8D%E3%83%B3%E3%82%B7%E3%83%B3%E3%82%BF%E3%83%BC%E3%82%BC&action=edit&redlink=1" class="new" title="「Δ-カジネンシンターゼ」 (存在しないページ)">δ-カジネンシンターゼ</a>（<a href="https://en.wikipedia.org/wiki/Delta-cadinene_synthase" class="extiw" title="en:Delta-cadinene synthase">英語版</a>）を減少させるためにRNAiが利用されている<a href="#cite_note-Sunilkumar-159">[159]</a>。同様に、<a href="/wiki/%E3%82%AD%E3%83%A3%E3%83%83%E3%82%B5%E3%83%90" title="キャッサバ">キャッサバ</a>で<a href="/wiki/%E3%82%B7%E3%82%A2%E3%83%B3%E5%8C%96%E7%89%A9" title="シアン化物">シアン化物</a>の原料となる<a href="/wiki/%E3%83%AA%E3%83%8A%E3%83%9E%E3%83%AA%E3%83%B3" title="リナマリン">リナマリン</a>を減少させる試みも行われている<a href="#cite_note-Siritunga-160">[160]</a>。 トマト類ではアレルゲンの減少<a href="#cite_note-Le_tomato-161">[161]</a>や<a href="/wiki/%E6%8A%97%E9%85%B8%E5%8C%96%E7%89%A9%E8%B3%AA" title="抗酸化物質">抗酸化物質</a>の強化<a href="#cite_note-Niggeweg-162">[162]</a>に成功している。<a href="/wiki/Flavr_Savr" title="Flavr Savr">Flavr Savr</a>トマトや<a href="/w/index.php?title=%E3%83%91%E3%83%91%E3%82%A4%E3%83%A4%E3%83%AA%E3%83%B3%E3%82%B0%E3%82%B9%E3%83%9D%E3%83%83%E3%83%88%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9&action=edit&redlink=1" class="new" title="「パパイヤリングスポットウイルス」 (存在しないページ)">パパイヤリングスポットウイルス</a>（<a href="https://en.wikipedia.org/wiki/Papaya_ringspot_virus" class="extiw" title="en:Papaya ringspot virus">英語版</a>）耐性<a href="/wiki/%E3%83%91%E3%83%91%E3%82%A4%E3%82%A2" title="パパイア">パパイヤ</a>の2品種など、これまで商業化されている品種はもともと<a href="/wiki/%E3%82%A2%E3%83%B3%E3%83%81%E3%82%BB%E3%83%B3%E3%82%B9RNA" title="アンチセンスRNA">アンチセンス</a>技術を用いて開発されたものであるが、実際にはRNAi経路が利用されている可能性が高い<a href="#cite_note-Sanders-163">[163]</a><a href="#cite_note-Chiang-164">[164]</a>。<a href="/w/index.php?title=%E3%82%A2%E3%82%B9%E3%83%9A%E3%83%AB%E3%82%AE%E3%83%AB%E3%82%B9%E3%83%BB%E3%83%95%E3%83%A9%E3%83%90%E3%82%B9&action=edit&redlink=1" class="new" title="「アスペルギルス・フラバス」 (存在しないページ)">アスペルギルス・フラバス</a>（<a href="https://en.wikipedia.org/wiki/Aspergillus_flavus" class="extiw" title="en:Aspergillus flavus">英語版</a>）Aspergillus flavusの<a href="/wiki/%CE%91-%E3%82%A2%E3%83%9F%E3%83%A9%E3%83%BC%E3%82%BC" class="mw-redirect" title="Α-アミラーゼ">α-アミラーゼ</a>（<a href="https://en.wikipedia.org/wiki/Alpha-amylase" class="extiw" title="en:Alpha-amylase">英語版</a>）を標的としたRNAiによるサイレンシングは<a href="/wiki/%E3%83%88%E3%82%A6%E3%83%A2%E3%83%AD%E3%82%B3%E3%82%B7" title="トウモロコシ">トウモロコシ</a>内でのこの菌類の増殖を低下せ、穀物を危険な<a href="/wiki/%E3%82%A2%E3%83%95%E3%83%A9%E3%83%88%E3%82%AD%E3%82%B7%E3%83%B3" title="アフラトキシン">アフラトキシン</a>による汚染から防ぐために利用されている<a href="#cite_note-:50-165">[165]</a>。<a href="/wiki/%E3%82%BF%E3%83%9E%E3%83%8D%E3%82%AE" title="タマネギ">タマネギ</a>での催涙因子合成酵素（lachrymatory factor synthase）のサイレンシングは切っても<a href="/wiki/%E6%B6%99%E3%81%AE%E5%87%BA%E3%81%AA%E3%81%84%E3%82%BF%E3%83%9E%E3%83%8D%E3%82%AE" title="涙の出ないタマネギ">涙の出ないタマネギ</a>の生産に、<a href="/wiki/%E3%82%BB%E3%82%A4%E3%83%A8%E3%82%A6%E3%82%A2%E3%83%96%E3%83%A9%E3%83%8A" title="セイヨウアブラナ">アブラナ</a>でのBP1遺伝子のサイレンシングは<a href="/wiki/%E5%85%89%E5%90%88%E6%88%90" title="光合成">光合成</a>の改善に利用されている<a href="#cite_note-:51-166">[166]</a>。<a href="/wiki/%E3%82%B3%E3%83%A0%E3%82%AE" title="コムギ">コムギ</a>では、<a href="/wiki/%E3%82%A2%E3%83%9F%E3%83%AD%E3%83%BC%E3%82%B9" title="アミロース">アミロース</a>含量の増大を目的としてSBEIIa、SBEIIb遺伝子が標的となっているほか<a href="#cite_note-:52-167">[167]</a>、六倍体品種の<a href="/wiki/%E6%A9%9F%E8%83%BD%E3%82%B2%E3%83%8E%E3%83%9F%E3%82%AF%E3%82%B9" title="機能ゲノミクス">機能ゲノミクス</a>研究にRNAiが、そしてLr21遺伝子によってもたらされる<a href="/w/index.php?title=%E3%82%B3%E3%83%A0%E3%82%AE%E8%B5%A4%E3%81%95%E3%81%B3%E7%97%85&action=edit&redlink=1" class="new" title="「コムギ赤さび病」 (存在しないページ)">コムギ赤さび病</a>（<a href="https://en.wikipedia.org/wiki/Wheat_leaf_rust" class="extiw" title="en:Wheat leaf rust">英語版</a>）耐性機構の研究のためにvirus-induced gene silencing（VIGS、RNAiの一種）が利用されている<a href="#cite_note-Downie-et-al-2021-168">[168]</a>。　 <div class="mw-heading mw-heading4"><h4 id="その他の作物">その他の作物</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=26" title="節を編集: その他の作物">編集</a>]</div> <a href="/wiki/%E3%82%BF%E3%83%90%E3%82%B3" title="タバコ">タバコ</a>では<a href="/wiki/%E7%99%BA%E7%99%8C%E6%80%A7" title="発癌性">発がん性</a>を有する可能性が高い物質の前駆体を減少させる取り組みが行われている<a href="#cite_note-Gavilano-169">[169]</a>。また実験室レベルでは、一般的な植物ウイルスに対する耐性の付与などの改変が行われている<a href="#cite_note-Zadeh-170">[170]</a>。<a href="/wiki/%E3%82%B1%E3%82%B7" title="ケシ">ケシ</a>による非<a href="/wiki/%E9%BA%BB%E8%96%AC" title="麻薬">麻薬</a>性<a href="/wiki/%E3%82%A2%E3%83%AB%E3%82%AB%E3%83%AD%E3%82%A4%E3%83%89" title="アルカロイド">アルカロイド</a>の産生も試みられている<a href="#cite_note-Allen-171">[171]</a>。 <div class="mw-heading mw-heading4"><h4 id="殺虫剤">殺虫剤</h4>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=27" title="節を編集: 殺虫剤">編集</a>]</div> <a href="/wiki/%E6%AE%BA%E8%99%AB%E5%89%A4" title="殺虫剤">殺虫剤</a>としてのRNAiの開発が行われており、遺伝子操作や外部からの投与など複数のアプローチがとられている<a href="#cite_note-s1308-2">[2]</a>。一部の昆虫の中腸の細胞は、environmental RNAiと呼ばれる過程でdsRNA分子を取り込むことがある<a href="#cite_note-:53-172">[172]</a>。一部の昆虫では、その効果は昆虫の体中に広がり、全身に作用する<a href="#cite_note-:54-173">[173]</a>。ヒトがこうした殺虫性のdsRNAを発現する遺伝子組み換え作物を消費することで予想される曝露量の数百万倍の量を曝露した場合でも、動物に悪影響はみられない<a href="#cite_note-:55-174">[174]</a>。 RNAiの影響は<a href="/wiki/%E9%B1%97%E7%BF%85%E7%9B%AE" class="mw-redirect" title="鱗翅目">鱗翅目</a>（チョウやガ）の生物種によってさまざまに異なり、おそらくそれは唾液や消化液のRNA分解能力の違いによるものである。cotton bollworm、<a href="/w/index.php?title=%E3%82%B7%E3%83%AD%E3%82%A4%E3%83%81%E3%83%A2%E3%82%B8%E3%83%A8%E3%83%88%E3%82%A6&action=edit&redlink=1" class="new" title="「シロイチモジヨトウ」 (存在しないページ)">シロイチモジヨトウ</a>（<a href="https://en.wikipedia.org/wiki/Beet_armyworm" class="extiw" title="en:Beet armyworm">英語版</a>）、<a href="/w/index.php?title=%E3%83%8B%E3%82%AB%E3%83%A1%E3%82%A4%E3%82%AC&action=edit&redlink=1" class="new" title="「ニカメイガ」 (存在しないページ)">ニカメイガ</a>（<a href="https://en.wikipedia.org/wiki/Chilo_suppressalis" class="extiw" title="en:Chilo suppressalis">英語版</a>）では給餌によるRNAi感受性は示されていない<a href="#cite_note-s1308-2">[2]</a>。 RNAiに対する耐性は広域的、すなわちある配列に対する耐性が他のdsRNA配列に対する耐性も付与する可能性が示唆されている。ある<a href="/w/index.php?title=%E3%82%A6%E3%82%A8%E3%82%B9%E3%82%BF%E3%83%B3%E3%82%B3%E3%83%BC%E3%83%B3%E3%83%AB%E3%83%BC%E3%83%88%E3%83%AF%E3%83%BC%E3%83%A0&action=edit&redlink=1" class="new" title="「ウエスタンコーンルートワーム」 (存在しないページ)">ウエスタンコーンルートワーム</a>（<a href="https://en.wikipedia.org/wiki/Western_corn_rootworm" class="extiw" title="en:Western corn rootworm">英語版</a>）の実験室集団では、腸からのDvSnf7を標的としたdsRNAの取り込みが起こらないために耐性が生じている<a href="#cite_note-:56-175">[175]</a>。DvSnf7に対する他のdsRNA配列を試した際にも有効性は見られず、耐性管理は単純にdsRNAの配列を切り替えるだけでは困難であることが示唆される。<a href="/wiki/%E3%83%90%E3%83%81%E3%83%AB%E3%82%B9%E3%83%BB%E3%83%81%E3%83%A5%E3%83%BC%E3%83%AA%E3%83%B3%E3%82%B2%E3%83%B3%E3%82%B7%E3%82%B9" title="バチルス・チューリンゲンシス">バチルス・チューリンゲンシス</a>Bacillus thuringiensis由来のCryタンパク質とRNAiなど複数の戦略を併用することで、耐性の出現は遅らせることができると考えられている<a href="#cite_note-s1308-2">[2]</a><a href="#cite_note-:57-176">[176]</a>。 <a href="/wiki/%E3%82%B7%E3%83%A7%E3%82%A6%E3%82%B8%E3%83%A7%E3%82%A6%E3%83%90%E3%82%A8%E5%B1%9E" class="mw-redirect" title="ショウジョウバエ属">ショウジョウバエ属</a>（<a href="https://en.wikipedia.org/wiki/Drosophila" class="extiw" title="en:Drosophila">英語版</a>）Drosophila spp.、<a href="/wiki/%E3%82%AB%E3%82%A4%E3%82%B3%E3%82%AC" class="mw-redirect" title="カイコガ">カイコガ</a>Bombyx mori、<a href="/wiki/%E3%83%88%E3%83%8E%E3%82%B5%E3%83%9E%E3%83%90%E3%83%83%E3%82%BF" title="トノサマバッタ">トノサマバッタ</a>属Locusta spp、<a href="/w/index.php?title=%E3%82%B9%E3%83%9D%E3%83%89%E3%83%97%E3%83%86%E3%83%A9%E5%B1%9E&action=edit&redlink=1" class="new" title="「スポドプテラ属」 (存在しないページ)">スポドプテラ属</a>（<a href="https://en.wikipedia.org/wiki/Spodoptera" class="extiw" title="en:Spodoptera">英語版</a>）Spodoptera spp.、<a href="/wiki/%E3%82%B3%E3%82%AF%E3%83%8C%E3%82%B9%E3%83%88%E3%83%A2%E3%83%89%E3%82%AD" title="コクヌストモドキ">コクヌストモドキ</a>Tribolium castaneum、<a href="/w/index.php?title=%E3%83%88%E3%83%93%E3%82%A4%E3%83%AD%E3%82%A6%E3%83%B3%E3%82%AB&action=edit&redlink=1" class="new" title="「トビイロウンカ」 (存在しないページ)">トビイロウンカ</a>（<a href="https://en.wikipedia.org/wiki/Brown_planthopper" class="extiw" title="en:Brown planthopper">英語版</a>）Nilaparvata lugens、<a href="/w/index.php?title=%E3%82%AA%E3%82%AA%E3%82%BF%E3%83%90%E3%82%B3%E3%82%AC&action=edit&redlink=1" class="new" title="「オオタバコガ」 (存在しないページ)">オオタバコガ</a>（<a href="https://en.wikipedia.org/wiki/Helicoverpa_armigera" class="extiw" title="en:Helicoverpa armigera">英語版</a>）Helicoverpa armigera、<a href="/wiki/%E3%82%BB%E3%82%A4%E3%83%A8%E3%82%A6%E3%83%9F%E3%83%84%E3%83%90%E3%83%81" title="セイヨウミツバチ">セイヨウミツバチ</a>Apis melliferaは、昆虫の特定の系統内でRNAiがどのように機能するかを知るために広く利用されているモデルである。<a href="/wiki/%E3%82%A4%E3%82%A8%E3%83%90%E3%82%A8" title="イエバエ">イエバエ</a>Musca domesticaはAgo2遺伝子を2つ持ち、<a href="/wiki/%E3%83%84%E3%82%A7%E3%83%84%E3%82%A7%E3%83%90%E3%82%A8" title="ツェツェバエ">ツェツェバエ</a>Glossina morsitansは3つ持つことが知られている<a href="#cite_note-Mongelli-Saleh-2016-177">[177]</a><a href="#cite_note-Zhu-Palli-2020-178">[178]</a>。miRNA経路に関しては、<a href="/w/index.php?title=%E3%83%AD%E3%82%B7%E3%82%A2%E3%82%B3%E3%83%A0%E3%82%AE%E3%82%A2%E3%83%96%E3%83%A9%E3%83%A0%E3%82%B7&action=edit&redlink=1" class="new" title="「ロシアコムギアブラムシ」 (存在しないページ)">ロシアコムギアブラムシ</a>（<a href="https://en.wikipedia.org/wiki/Russian_wheat_aphid" class="extiw" title="en:Russian wheat aphid">英語版</a>）Diuraphis noxiaは2つのAgo1、M. domesticaは2つのDcr1、<a href="/w/index.php?title=%E3%82%A8%E3%83%B3%E3%83%89%E3%82%A6%E3%83%92%E3%82%B2%E3%83%8A%E3%82%AC%E3%82%A2%E3%83%96%E3%83%A9%E3%83%A0%E3%82%B7&action=edit&redlink=1" class="new" title="「エンドウヒゲナガアブラムシ」 (存在しないページ)">エンドウヒゲナガアブラムシ</a>（<a href="https://en.wikipedia.org/wiki/Acyrthosiphon_pisum" class="extiw" title="en:Acyrthosiphon pisum">英語版</a>）Acyrthosiphon pisumはAgo1、Loqs、Dcr1を2つずつ持ち、Pashaを4つ持つ。<a href="/w/index.php?title=PiRNA&action=edit&redlink=1" class="new" title="「PiRNA」 (存在しないページ)">piRNA</a>（<a href="https://en.wikipedia.org/wiki/Piwi-interacting_RNA" class="extiw" title="en:Piwi-interacting RNA">英語版</a>）に関しては、G. morsitansとA. pisumは2つまたは3つのAgo3を持つ<a href="#cite_note-Zhu-Palli-2020-178">[178]</a>。こうした研究により、将来的な殺虫剤開発の標的や、作用機序、他の殺虫剤に対する耐性の理由などが明らかとなった<a href="#cite_note-Zhu-Palli-2020-178">[178]</a>。 <div class="mw-heading mw-heading5"><h5 id="トランスジェニック植物">トランスジェニック植物</h5>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=28" title="節を編集: トランスジェニック植物">編集</a>]</div> トランスジェニック作物はdsRNAを発現するように作製されており、その配列は標的害虫の重要な遺伝子をサイレンシングするよう慎重に選ばれたものである。こうしたdsRNAは、特定の遺伝子配列を発現する昆虫のみに影響を与えるよう設計されている。2009年の実証実験では、RNAが4種のショウジョウバエのうちいずれか1種のみに対して殺虫作用を示し、他の3種には害を及ぼさないことが示された<a href="#cite_note-s1308-2">[2]</a>。 2012年、<a href="/wiki/%E3%82%B7%E3%83%B3%E3%82%B8%E3%82%A7%E3%83%B3%E3%82%BF" title="シンジェンタ">シンジェンタ</a>はベルギーのRNAi企業Devgenを5億2200万ドルで買収し、<a href="/wiki/%E3%83%A2%E3%83%B3%E3%82%B5%E3%83%B3%E3%83%88_(%E4%BC%81%E6%A5%AD)" title="モンサント (企業)">モンサント</a>は<a href="/w/index.php?title=%E3%82%A2%E3%83%AB%E3%83%8A%E3%82%A4%E3%83%A9%E3%83%A0%E3%83%BB%E3%83%95%E3%82%A1%E3%83%BC%E3%83%9E%E3%82%B7%E3%83%A5%E3%83%BC%E3%83%86%E3%82%A3%E3%82%AB%E3%83%AB%E3%82%BA&action=edit&redlink=1" class="new" title="「アルナイラム・ファーマシューティカルズ」 (存在しないページ)">アルナイラム・ファーマシューティカルズ</a>（<a href="https://en.wikipedia.org/wiki/Alnylam_Pharmaceuticals" class="extiw" title="en:Alnylam Pharmaceuticals">英語版</a>）から<a href="/wiki/%E7%9F%A5%E7%9A%84%E8%B2%A1%E7%94%A3%E6%A8%A9" title="知的財産権">知的財産権</a>の独占権を2920万ドルで取得した。ペルー・リマの<a href="/w/index.php?title=%E5%9B%BD%E9%9A%9B%E3%83%9D%E3%83%86%E3%83%88%E3%82%BB%E3%83%B3%E3%82%BF%E3%83%BC&action=edit&redlink=1" class="new" title="「国際ポテトセンター」 (存在しないページ)">国際ポテトセンター</a>（<a href="https://en.wikipedia.org/wiki/International_Potato_Center" class="extiw" title="en:International Potato Center">英語版</a>）では、幼虫による<a href="/wiki/%E3%82%B5%E3%83%84%E3%83%9E%E3%82%A4%E3%83%A2" title="サツマイモ">サツマイモ</a>の食害が世界的に問題となっている<a href="/wiki/%E3%82%A2%E3%83%AA%E3%83%A2%E3%83%89%E3%82%AD%E3%82%BE%E3%82%A6%E3%83%A0%E3%82%B7" title="アリモドキゾウムシ">アリモドキゾウムシ</a>に対する標的遺伝子の探索が行われている。他にも、アリ、毛虫、pollen beetleなどの遺伝子のサイレンシングが試みられている。モンサントは、アメリカ合衆国だけで毎年10億ドルの被害をもたらしているウエスタンコーンルートワームのSnf7遺伝子を標的としたdsRNAを発現する、トランスジェニックトウモロコシの種子を初めて販売することとなる可能性が高い。2012年の論文では、Snf7のサイレンシングは幼虫の成長を止め、数日以内に死滅させることが示されている。2013年に同チームは、このRNAが他の生物種に影響を与えることはほとんどないことを示した<a href="#cite_note-s1308-2">[2]</a>。 <div class="mw-heading mw-heading5"><h5 id="外部からの投与">外部からの投与</h5>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=29" title="節を編集: 外部からの投与">編集</a>]</div> dsRNAは遺伝子組み換え以外の方法でも供給することができる。1つのアプローチは、<a href="/wiki/%E7%81%8C%E6%BC%91" title="灌漑">灌漑</a>用水への添加である。RNA分子は植物の<a href="/wiki/%E7%B6%AD%E7%AE%A1%E6%9D%9F" title="維管束">維管束</a>系へ吸収され、その植物を食べる昆虫を殺す。他のアプローチは、従来の農薬のような形でのdsRNAの噴霧である。こうした方法は耐性の出現に対してより早く対応することができるが、dsRNAの低コストでの作製法を必要とし、そうした手法は現在のところ存在しない<a href="#cite_note-s1308-2">[2]</a>。 <div class="mw-heading mw-heading3"><h3 id="ゲノムスケールでのスクリーニング">ゲノムスケールでのスクリーニング</h3>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=30" title="節を編集: ゲノムスケールでのスクリーニング">編集</a>]</div> ゲノムスケールでのRNAi研究は、ハイスループットスクリーニング（HTS）技術に依存している。RNAi HTS技術はゲノムワイドな機能喪失スクリーニングを可能にし、特定の表現型と関係する遺伝子の同定に広く利用されている。この技術は、遺伝子発現<a href="/wiki/%E3%83%9E%E3%82%A4%E3%82%AF%E3%83%AD%E3%82%A2%E3%83%AC%E3%82%A4" title="マイクロアレイ">マイクロアレイ</a>や<a href="/wiki/%E4%B8%80%E5%A1%A9%E5%9F%BA%E5%A4%9A%E5%9E%8B" title="一塩基多型">一塩基多型</a>発見プラットフォームといった<a href="/wiki/%E3%82%B2%E3%83%8E%E3%83%9F%E3%82%AF%E3%82%B9" title="ゲノミクス">ゲノミクス</a>の第一の波に続く、第二の波となる可能性があるとの評価がなされている<a href="#cite_note-Matson-179">[179]</a>。ゲノムスケールでのRNAiスクリーニングの大きな利点の1つは、数千もの遺伝子を同時に調査することができる点である。ゲノムスケールでのRNAiスクリーニングでは1つの実験から大量のデータが生み出されるため、データ生成量の爆発的増加をもたらしている。こうした巨大なデータセットの処理は基本的な課題となっており、適切な統計学やバイオインフォマティクス的手法を必要とする。細胞ベースのRNAiスクリーニングの基本的過程は、RNAiライブラリや頑強で安定した細胞種の選択、RNAi試薬によるトランスフェクション、処理とインキュベーション、シグナル検出、重要な遺伝子または治療標的遺伝子の解析、同定などからなる<a href="#cite_note-ZhangXHD-180">[180]</a>。 <div class="mw-heading mw-heading2"><h2 id="歴史">歴史</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=31" title="節を編集: 歴史">編集</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/%E3%83%95%E3%82%A1%E3%82%A4%E3%83%AB:Rnai_phenotype_petunia_crop.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_phenotype_petunia_crop.png/250px-Rnai_phenotype_petunia_crop.png" decoding="async" width="250" height="70" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_phenotype_petunia_crop.png/375px-Rnai_phenotype_petunia_crop.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/24/Rnai_phenotype_petunia_crop.png/500px-Rnai_phenotype_petunia_crop.png 2x" data-file-width="1363" data-file-height="382" /></a><figcaption>RNAiによって色素形成のための遺伝子がサイレンシングされたペチュニアの例。左は野生型、右の2つでは導入遺伝子と内在性遺伝子の双方の発現の抑制を誘導する遺伝子導入の結果、色素を持たない白い領域が生じている<a href="#cite_note-Matzke-181">[181]</a>。</figcaption></figure> RNAi過程は、それがRNAと関連した機構であることが知られる前には、"co-suppression"や"quelling"と呼ばれていた。RNAiの発見に先立って、トランスジェニック植物で発現させたアンチセンスRNAによる転写阻害がまず観察され<a href="#cite_note-Ecker_1986-182">[182]</a>、そして1990年代初頭のアメリカ合衆国とオランダの植物学者によって行われた実験による予想外の結果の報告<a href="#cite_note-Napoli_1990-183">[183]</a>によってより直接的にRNAiの発見への道が開かれた。この実験では<a href="/wiki/%E3%83%9A%E3%83%81%E3%83%A5%E3%83%8B%E3%82%A2" title="ペチュニア">ペチュニア</a>の花の色の変化が試みられ、研究者らは正常なピンクまたはスミレ色の花のペチュニアに対し、花の色素形成に重要な酵素である<a href="/w/index.php?title=%E3%82%AB%E3%83%AB%E3%82%B3%E3%83%B3%E3%82%B7%E3%83%B3%E3%82%BF%E3%83%BC%E3%82%BC&action=edit&redlink=1" class="new" title="「カルコンシンターゼ」 (存在しないページ)">カルコンシンターゼ</a>（<a href="https://en.wikipedia.org/wiki/Chalcone_synthase" class="extiw" title="en:Chalcone synthase">英語版</a>）をコードする遺伝子のさらなるコピーを導入した。コピー数の増加による遺伝子の過剰発現によってより濃い色の花となることが予想されたが、実際には一部の花では紫色の色素は薄くなり、そして斑入りのパターンが形成されることもあった。このことは、カルコンシンターゼの活性は状況依存的に大きく低下するか、または抑制されていることを示していた。後に、一部の形質転換体のゲノム中のさまざまな位置で反対向きのプロモーターに隣接して導入遺伝子が挿入された結果、プロモーターの活性化によってアンチセンス転写産物が発現し、遺伝子がサイレンシングされたという説明がなされた。初期のRNAiの観察の他の例としては<a href="/wiki/%E3%82%A2%E3%82%AB%E3%83%91%E3%83%B3%E3%82%AB%E3%83%93" title="アカパンカビ">アカパンカビ</a>Neurospora crassaの研究のものがあるが<a href="#cite_note-Romano-184">[184]</a>、これが関連した現象であるとはすぐには認識されなかった。植物での現象のさらなる研究によって、ダウンレギュレーションはmRNAの分解率の上昇を介した、遺伝子発現の転写後阻害によるものであることが示された<a href="#cite_note-Van_Blokland_1994-185">[185]</a>。この現象は"co-suppression of gene expression"と呼ばれたが、その分子機構はいまだ不明のままであった<a href="#cite_note-:58-186">[186]</a>。 それから間もなく、ウイルス病に対する植物の耐性の改善に取り組んでいた植物ウイルス学者らによって、類似した予想外の現象が観察された。ウイルス特異的タンパク質を発現する植物はウイルス感染に対するトレランス（tolerance）や抵抗性（resistance）の向上がみられることは知られていたが、ウイルスRNA配列の短い非コード領域のみを持つ植物も同様の防御レベルを示すという予想外の結果が得られた。研究者らは導入遺伝子によって産生されるウイルスRNAがウイルスの複製を阻害すると考えた<a href="#cite_note-Covey_1997-187">[187]</a>。逆実験として、植物遺伝子の短い配列を導入したウイルスは、感染した植物で標的遺伝子を抑制することが示された<a href="#cite_note-Kumagai_1995-188">[188]</a>。この現象は"virus-induced gene silencing"（VIGS）と呼ばれ、これらの現象はまとめて"post transcriptional gene silencing"と呼ばれるようになった<a href="#cite_note-Ratcliff_1997-189">[189]</a>。 こうした植物における初期の観察の後、他の生物種におけるこうした現象の探索が行われた<a href="#cite_note-Guo-190">[190]</a><a href="#cite_note-Pal-Bhadra-191">[191]</a>。クレイグ・メローとアンドリュー・ファイアーによる1998年のNature誌の論文では、C. elegansに二本鎖RNAを注入した後に強力な遺伝子サイレンシング効果がみられることが報告された<a href="#cite_note-Fire-192">[192]</a>。彼らは筋タンパク質の産生の調節の研究の際に、mRNAやアンチセンスRNAの注入はタンパク質産生に影響を及ぼさないが、二本鎖RNAの注入によって標的遺伝子がサイレンシングされることを発見した。この研究をもとに、RNAiという用語を作った。この発見は、この現象の原因となる因子を初めて同定したこととなる。ファイアーとメローは2006年にノーベル生理学・医学賞を受賞した<a href="#cite_note-Daneholt2006-34">[34]</a>。 <div class="mw-heading mw-heading2"><h2 id="出典">出典</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=32" title="節を編集: 出典">編集</a>]</div> <div class="reflist" style="-moz-column-count:2; -webkit-column-count:2; column-count:2; -moz-column-width: 20em; -webkit-column-width: 20em; column-width: 20em; list-style-type: decimal;"> <ol class="references"> <li id="cite_note-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564-1">^ <a href="#cite_ref-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564_1-0">a</a> <a href="#cite_ref-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564_1-1">b</a> <a href="#cite_ref-Saurabh_Satyajit,_Vidyarthi_AS,_Prasad_D_2014_543–564_1-2">c</a> <style 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Cambridge University Press. pp. ix–xiii. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r101121245"><a href="/wiki/ISBN" title="ISBN">ISBN</a> <a href="/wiki/%E7%89%B9%E5%88%A5:%E6%96%87%E7%8C%AE%E8%B3%87%E6%96%99/978-0-521-73444-8" title="特別:文献資料/978-0-521-73444-8">978-0-521-73444-8</a>. <a rel="nofollow" class="external free" href="http://www.cambridge.org/9780521734448">http://www.cambridge.org/9780521734448</a></cite><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Optimal+High-Throughput+Screening%3A+Practical+Experimental+Design+and+Data+Analysis+for+Genome-scale+RNAi+Research&rft.aulast=Zhang+XHD&rft.au=Zhang+XHD&rft.date=2011&rft.pages=pp.%26nbsp%3Bix%E2%80%93xiii&rft.pub=Cambridge+University+Press&rft.isbn=978-0-521-73444-8&rft_id=http%3A%2F%2Fwww.cambridge.org%2F9780521734448&rfr_id=info:sid/ja.wikipedia.org:RNA%E5%B9%B2%E6%B8%89">  </li> <li id="cite_note-Matzke-181"><a href="#cite_ref-Matzke_181-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r101121245"><cite id="CITEREFMatzkeMatzke2004" class="citation journal cs1">Matzke MA, Matzke AJ (2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC406394">"Planting the Seeds of a New Paradigm"</a>. 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"A similarity between viral defense and gene silencing in plants". Science. 276 (5318): 1558–60. <a href="/wiki/Doi_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Doi (識別子)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.276.5318.1558">10.1126/science.276.5318.1558</a>. <a href="/wiki/PMID_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="PMID (識別子)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18610513">18610513</a>。</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=A+similarity+between+viral+defense+and+gene+silencing+in+plants&rft.volume=276&rft.issue=5318&rft.pages=1558-60&rft.date=1997-06&rft_id=info%3Adoi%2F10.1126%2Fscience.276.5318.1558&rft_id=info%3Apmid%2F18610513&rft.aulast=Ratcliff&rft.aufirst=F&rft.au=Harrison%2C+BD&rft.au=Baulcombe%2C+DC&rfr_id=info%3Asid%2Fja.wikipedia.org%3ARNA%E5%B9%B2%E6%B8%89" class="Z3988"> </li> <li id="cite_note-Guo-190"><a href="#cite_ref-Guo_190-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r101121245"><cite id="CITEREFGuoKemphues1995" class="citation journal cs1">Guo S, Kemphues KJ (May 1995). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F0092-8674%2895%2990082-9">"par-1, a gene required for establishing polarity in C. elegans embryos, encodes a putative Ser/Thr kinase that is asymmetrically distributed"</a>. Cell. 81 (4): 611–20. <a href="/wiki/Doi_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Doi (識別子)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F0092-8674%2895%2990082-9">10.1016/0092-8674(95)90082-9</a>. <a href="/wiki/PMID_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="PMID (識別子)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/7758115">7758115</a>。</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=par-1%2C+a+gene+required+for+establishing+polarity+in+C.+elegans+embryos%2C+encodes+a+putative+Ser%2FThr+kinase+that+is+asymmetrically+distributed&rft.volume=81&rft.issue=4&rft.pages=611-20&rft.date=1995-05&rft_id=info%3Adoi%2F10.1016%2F0092-8674%2895%2990082-9&rft_id=info%3Apmid%2F7758115&rft.aulast=Guo&rft.aufirst=S&rft.au=Kemphues%2C+KJ&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252F0092-8674%252895%252990082-9&rfr_id=info%3Asid%2Fja.wikipedia.org%3ARNA%E5%B9%B2%E6%B8%89" class="Z3988"> </li> <li id="cite_note-Pal-Bhadra-191"><a href="#cite_ref-Pal-Bhadra_191-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r101121245"><cite id="CITEREFPal-BhadraBhadraBirchler1997" class="citation journal cs1">Pal-Bhadra M, Bhadra U, Birchler JA (August 1997). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0092-8674%2800%2980508-5">"Cosuppression in Drosophila: gene silencing of Alcohol dehydrogenase by white-Adh transgenes is Polycomb dependent"</a>. Cell. 90 (3): 479–90. <a href="/wiki/Doi_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Doi (識別子)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0092-8674%2800%2980508-5">10.1016/S0092-8674(00)80508-5</a>. <a href="/wiki/PMID_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="PMID (識別子)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9267028">9267028</a>。</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=Cosuppression+in+Drosophila%3A+gene+silencing+of+Alcohol+dehydrogenase+by+white-Adh+transgenes+is+Polycomb+dependent&rft.volume=90&rft.issue=3&rft.pages=479-90&rft.date=1997-08&rft_id=info%3Adoi%2F10.1016%2FS0092-8674%2800%2980508-5&rft_id=info%3Apmid%2F9267028&rft.aulast=Pal-Bhadra&rft.aufirst=M&rft.au=Bhadra%2C+U&rft.au=Birchler%2C+JA&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252FS0092-8674%252800%252980508-5&rfr_id=info%3Asid%2Fja.wikipedia.org%3ARNA%E5%B9%B2%E6%B8%89" class="Z3988"> </li> <li id="cite_note-Fire-192"><a href="#cite_ref-Fire_192-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r101121245"><cite id="CITEREFFireXuMontgomeryKostas1998" class="citation journal cs1">Fire A, Xu S, Montgomery MK, Kostas SA, Driver SE, Mello CC (February 1998). "Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans". Nature. 391 (6669): 806–11. <a href="/wiki/Bibcode_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Bibcode (識別子)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1998Natur.391..806F">1998Natur.391..806F</a>. <a href="/wiki/Doi_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Doi (識別子)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F35888">10.1038/35888</a>. <a href="/wiki/PMID_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="PMID (識別子)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9486653">9486653</a>. <a href="/wiki/S2CID_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="S2CID (識別子)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4355692">4355692</a>。</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Potent+and+specific+genetic+interference+by+double-stranded+RNA+in+Caenorhabditis+elegans&rft.volume=391&rft.issue=6669&rft.pages=806-11&rft.date=1998-02&rft_id=info%3Adoi%2F10.1038%2F35888&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4355692%23id-name%3DS2CID&rft_id=info%3Apmid%2F9486653&rft_id=info%3Abibcode%2F1998Natur.391..806F&rft.aulast=Fire&rft.aufirst=A&rft.au=Xu%2C+S&rft.au=Montgomery%2C+MK&rft.au=Kostas%2C+SA&rft.au=Driver%2C+SE&rft.au=Mello%2C+CC&rfr_id=info%3Asid%2Fja.wikipedia.org%3ARNA%E5%B9%B2%E6%B8%89" class="Z3988"> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="外部リンク">外部リンク</h2>[<a href="/w/index.php?title=RNA%E5%B9%B2%E6%B8%89&action=edit&section=33" title="節を編集: 外部リンク">編集</a>]</div> <ul><li><a rel="nofollow" class="external text" href="http://www.thenakedscientists.com/HTML/articles/article/rna-interference-explained/">Overview of the RNAi process</a>, from Cambridge University's The Naked Scientists</li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/focus/rnai/animations/animation/animation.htm">Animation of the RNAi process</a>, from <a href="/wiki/Nature_(journal)" class="mw-redirect" title="Nature (journal)">Nature</a></li> <li><a rel="nofollow" class="external text" href="https://www.pbs.org/wgbh/nova/sciencenow/3210/02.html">NOVA scienceNOW explains RNAi</a> – A 15-minute video of the Nova broadcast that aired on <a href="/wiki/Public_Broadcasting_Service" class="mw-redirect" title="Public Broadcasting Service">PBS</a>, 26 July 2005</li> <li><a rel="nofollow" class="external text" href="http://www.silencinggenomes.org/">Silencing Genomes</a> RNA interference (RNAi) experiments and bioinformatics in C. elegans for education. From the Dolan DNA Learning Center of Cold Spring Harbor Laboratory.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20061116000124/http://www.natureprotocols.com/2006/07/19/rnai_screens_in_c_elegans_in_a.php">RNAi screens in C. elegans in a 96-well liquid format and their application to the systematic identification of genetic interactions (a protocol)</a></li> <li><a rel="nofollow" class="external text" href="https://www.nytimes.com/2006/10/03/science/03nobel.html?ex=1180065600&en=e26ec371cb398ea0&ei=5070">2 American ‘Worm People’ Win Nobel for RNA Work</a>, from <a href="/wiki/The_New_York_Times" class="mw-redirect" title="The New York Times">NY Times</a></li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/mt/webfocus/rnai/index.html#2008">Molecular Therapy web focus: "The development of RNAi as a therapeutic strategy"</a>, a collection of free articles about RNAi as a therapeutic strategy.</li> <li><a rel="nofollow" class="external text" href="http://www.genomernai.de/GenomeRNAi/">GenomeRNAi</a>: a database of phenotypes from RNA interference screening experiments in Drosophila melanogaster and Homo sapians</li> <li><a rel="nofollow" class="external text" href="http://dharmacon.horizondiscovery.com/rnai-and-custom-rna-synthesis/">RNAi tools</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180619035904/http://dharmacon.horizondiscovery.com/rnai-and-custom-rna-synthesis/">Archived</a> 19 June 2018 at the <a href="/wiki/%E3%82%A6%E3%82%A7%E3%82%A4%E3%83%90%E3%83%83%E3%82%AF%E3%83%9E%E3%82%B7%E3%83%B3" title="ウェイバックマシン">Wayback Machine</a>. Pre-designed and custom RNA Interference tools</li> <li><cite style="font-style:normal" class="citation" id="CITEREF塩見2015"><a href="/wiki/%E5%A1%A9%E8%A6%8B%E7%BE%8E%E5%96%9C%E5%AD%90" title="塩見美喜子">塩見美喜子</a>『<a rel="nofollow" class="external text" href="https://doi.org/10.14931/bsd.3744">RNA干渉</a>』INCF Japan Node、2015年。<a href="/wiki/Doi_(%E8%AD%98%E5%88%A5%E5%AD%90)" class="mw-redirect" title="Doi (識別子)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.14931%2Fbsd.3744">10.14931/bsd.3744</a>。<a rel="nofollow" class="external free" href="https://doi.org/10.14931/bsd.3744">https://doi.org/10.14931/bsd.3744</a>。2022年4月17日閲覧。</cite><span class="Z3988" 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