JP5969889B2 - 二本鎖rnaによる遺伝子阻害 - Google Patents
二本鎖rnaによる遺伝子阻害 Download PDFInfo
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- JP5969889B2 JP5969889B2 JP2012229547A JP2012229547A JP5969889B2 JP 5969889 B2 JP5969889 B2 JP 5969889B2 JP 2012229547 A JP2012229547 A JP 2012229547A JP 2012229547 A JP2012229547 A JP 2012229547A JP 5969889 B2 JP5969889 B2 JP 5969889B2
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Description
本発明は、国立衛生研究所により与えられた、助成金番号GM−37706、GM−17164、HD−33769およびGM−07231による米国政府補助で行われた。米国政府は、本発明にある種の権利を有する。
1.発明の分野
本発明は、二本鎖リボ核酸(dsRNA)による遺伝子発現の遺伝子特異的阻害に関する。
遺伝子発現の選択的阻害はバイオテクノロジーおよび遺伝子工学における長期にわたる切実なニーズであった。このゴールを達成するために大きな努力が投じられてきたが、この問題に対するより総括的な解決が未だ必要とされている
アンチセンス技術は、遺伝子特異的な干渉を達成するためのプロトコルにおいて最も一般的に記載された方法であった。アンチセンス戦略として、目的とする遺伝子に対するメッセンジャーRNAに相補的な一本鎖核酸の化学量論的量を細胞に導入する。アンチセンスを基礎とする方法のいくつかの困難性は、運搬、安定性、および必要容量に関するものである。一般的に、細胞は一本鎖核酸に対する取り込み機構を有していない、それゆえ非修飾一本鎖物質の取り込みは極端に非効率である。細胞に取り込まれるのを待つ間に、一本鎖物質は分解してしまう。なぜなら、アンチセンス干渉には、干渉物質が比較的高濃度(内部mRNAの濃度またはそれ以上)で蓄積することが必要であり、運搬されるべき必要量が効率の主な妨げである。結果として、アンチセンス技術の開発において、多くの努力が、ヌクレアーゼ消化に対して安定であり、そして細胞に容易に拡散して行くことができる修飾核酸の製造に焦点を合わせてきた。遺伝子治療または他の全身への適用のためにアンチセンス干渉を使用することは、非天然類縁体からの合成に必要な大量のオリゴヌクレオチド、その合成費用、および各細胞に高投与量を以ってしても干渉物質を充分な濃度で均一なプールを維持することの困難さによって制限されてきた。
第二の、提案された遺伝子工学処理された干渉方法は、三重らせん核酸構造を基礎とするものである。この方法は、三本鎖構造をとるためのある種の核酸集団のまれな性能による。生理的条件下においては、核酸は事実上すべて一本鎖または二本鎖であり、そして三本鎖を形成するとしてもまれである。しかし、時々、ある種の単純なプリンまたはピリミジンに富む配列は、極端なpH条件下のin vitroで(即ち、試験管内において)三本鎖分子を形成することができることが知られている。そのような構造は、一般に生理条件下での非常に一過性のものであるから、三本鎖構造を作ることを意図した非修飾核酸の単純な運搬は、干渉を生じることはない。アンチセンスについても、in vivoで使用するための三本鎖技術の開発は、in vivoで細胞によってより安定でより容易に吸収される修飾核酸の開発に焦点が合わされてきた。この技術の開発の付加的目標は、三本鎖物質の形成が生理的pHにおいて効率的に進行するような修飾核酸を製造することであった。
遺伝子特異的な干渉の第三の方法は、「共抑制」という名称でまとめられた操作手順の一群である。この方法は、植物で最初記載され、非連鎖で相同遺伝子の沈黙の原因となる導入遺伝子の作用に関するものであった。より最近になって、共抑制に類似の現象が、二つの動物:C.elegansおよびDrosophila、で報告された。共抑制は最初、強力で有用な遺伝子座の過剰発現を達成する試みで導入遺伝子を用いたグループからの報告で、偶然に観察された。ある場合には、過剰発現は成功したが、しかし多くの場合、結果は予測とは逆であった。それらの場合、導入遺伝子植物は実際内部遺伝子のより弱い発現を示した。植物における導入遺伝子介在共抑制について、いくつかの機構の提案がなされたが、それら機構についての提案のすべては仮説として残り、プロセスの決定的な機構の記述は提案されていない。共抑制を説明するために提案されたモデルは二つの異なったカテゴリーに置くことができる。一組の提案においては、二つの異なった染色体位置の間のDNAまたはクロマチンレベルでの直接の物理的相互作用が起こると仮定し、未だ未知の機構が遺伝子発現のde novoメチル化および引続く抑制を導くとしている。別のものは、導入遺伝子座で合成されるある種のRNA中間体を想定し、それが内部遺伝子に干渉をもたらすよう作用するとしている。RNAに干渉する性質および干渉過程の本質は決定されていない。最近、RNAウイルスでの一連の実験で、干渉過程でのRNA中間体の可能性をいく分か支持することが提案された。これらの実験で、複製されるRNAウイルスは、目的とする遺伝子からの分節を含むよう修飾される。この修飾ウイルスは、次いで、内部遺伝子の発現に干渉する作用があるかを試験される。この技術の初めの結果は、勇気付けられるものであったが、しかし、干渉効果の原因であるウイルスRNAの性質は決定されておらず、そしていずれの場合においても、植物ウイルスの宿主である植物に限定されていた。
本発明は、方法および効果の両者において、アンチセンス介在干渉と異なっている。アンチセンス介在遺伝子干渉法は、内部mRNAの濃度と等しいか或いはより大きい濃度で特異的一本鎖核酸分子を細胞内部へ運搬するという、主な課題を有している。二本鎖RNA介在阻害は、運搬されるべき物質の安定性および効果的な阻害に必要な濃度の両方で、有利性を有している。以下に、我々は、モデル生物体C.elegansにおいて、本発明は同様のアンチセンス法よりも少なくとも100倍、より効果的である(即ち、dsRNAは、遺伝子発現を減少させることにおいて、精製アンチセンスRNAの注入よりも少なくとも100倍、より効果的である)ことを開示する。それらの比較は、また、二本鎖RNAによる阻害はアンチセンス干渉とは異なった機構により起こることを明らかにしている。
三本鎖形成についての限られたデータは、本発明の安定な三本鎖中間体の関与に対して異議を唱えている。三本鎖構造は、あったとしても、生理条件下でまれに起こり、プリンとピリミジンの非常にまれな長い塩基配列に限られている。対照的に、dsRNA介在阻害は生理条件下で効率よく起こり、広い種々の阻害と標的ヌクレオチド配列で起こる。本発明は、公知の機能を有する遺伝子で変異なしに表現型模写を起こし、18の異なった遺伝子の発現を阻害するのに使用された。三重らせん形成の極端な環境および配列の束縛は、C.elegansにおけるdsRNA介在阻害が三重鎖構造により媒介されるということを起こりそうもなくしている。
共抑制と呼ばれる標的遺伝子介在遺伝子干渉は、広い種々の異なった過程を含んでいる。別のタイプの生物体に応用する観点からみると、植物の共抑制現象を拡張するのは困難である。共抑制を基にした一般的技術を創出することにおいてややこしくなるのは、植物におけるある種の導入遺伝子は内部遺伝子座の抑制を導き、あるものはそうでないと云うことである。C.elegansおよびDrosophilaでの成績は、ある種の導入遺伝子は干渉の原因となる(即ち、対応する内部遺伝子座の活性が定量的に減少する)が、しかし殆どの導入遺伝子はそのような効果を生み出さないことを示している。植物、線虫、および昆虫における予測される効果の欠如は、遺伝子発現を阻害するためにゲノムに導入遺伝子を単純に加えることの有用性を大きく制限する。植物におけるウイルス介在共抑制は非常に効果的にみえるが、しかし多くの欠点を有している。第一に、ウイルス構造のどのような面が観察された干渉について臨界的かが明確でないことである。他の系に拡張するには、そのような性質を有する系で有用なウイルスを見つけ出すことを要し、そしてそのような有用なウイルス剤のライブラリーは多くの生物体では得ることができない。第二に、遺伝子変化に有効な、生物体内で複製するウイルスの使用(即ち、長期または短期の遺伝子治療)には、本発明において定義される核酸を使用するよりも、有害な効果に対してかなりモニタリングや監視を必要とする。
細胞内の標的遺伝子の発現を阻害するための方法が提供される。本方法は、部分的または完全な二本鎖の性状のRNAを、細胞内または細胞外環境に導入することを含む。阻害は、標的遺伝子の部分からのヌクレオチド配列が阻害性RNAを作るように選択されることで、特異的である。我々は、本方法が、(1)遺伝子発現の阻害を作り出すのに効果的であり、(2)標的遺伝子に特異的であり、そして(3)標的遺伝子の多くの異なった型の阻害が一般的にできる、ことを開示する。
本発明は、二本鎖RNA(dsRNA)を導入することによる遺伝子発現の配列特異的阻害を作り出す方法を提供する。細胞内の標的遺伝子の発現を阻害するための方法が提供される。本方法は、部分的または完全な二本鎖の性状のRNAを細胞内へ導入することを含む。阻害は配列特異的であり、標的遺伝子の一部分からのヌクレオチド配列が阻害性RNAを作るように選択されることで、配列特異的である。我々は、本方法が、(1)遺伝子発現の阻害を作り出すのに効果的であり、(2)標的遺伝子に特異的であり、そして(3)標的遺伝子の多くの異なった型の阻害が一般的にできる、ことを開示する。
本発明の操作を、モデル遺伝子生物体Caenorhabditis elegannsで示した。
unc−22遺伝子活性と動物の可動性表現型の間の半定量的比較を与える予備的遺伝子解析の結果から、活性の初期比較するため、unc−22遺伝子を選択した3、8。活性の減少は激しい痙攣表現型の増加をもたらしたが、完全な機能の喪失は筋肉構造の欠損および減少した自発運動性の付加的出現をもたらした。unc−22は、多くの、しかし非必須の筋肉線維タンパク質をコードしている7−9。unc−22mRNAは、横紋筋細胞当り数千コピー存在する3。
RNAをT3およびT7RNAポリメラーゼのファージミドクローンから合成し6、次いで二つの逐次DNase処理で鋳型の除去を行った。センス、アンチセンス、および混合RNA集団を比較しなければならない場合においては、RNAは、低ゲル化温度アガロースの電気泳動により精製した。ゲル精製産物は、元の「センス」および「アンチセンス」調製物に見られた少数のバンドの多くがなくなっていた。それでもなお、精製RNA調製物の10%以下のRNA種が観察されなかった。ゲル精製なしに、「センス」および「アンチセンス」調製物は著しい阻害を生じた。この阻害活性は、ゲル精製によって減少されたか除去された。対照的に、ゲル精製および非ゲル精製のRNA調製物のセンス+アンチセンス混合物は、同じ効果を生じた。
内部遺伝子の阻害は、一般的に野生型遺伝的背景(N2)で測定される。解析される特徴は、行動、食餌、ふ化、体型、性的同一性、および生殖能力であった。gfp27およびlacZ活性の阻害はPD4251株を用いて評価された。この株は、三つのプラスミド:pSAK4(ミトコンドリア標的GFPを誘導するmyo−3プロモーター)、pSAK2(核標的GFP−LacZ融合を誘導するmyo−3プロモーター)、および選択し得るマーカーとしてdpy−20サブクローン27、より作られた集積アレイ(ccIs4251)を含有する安定したトランスジェニック株である。この株は、全ての体筋肉で、ミトコンドリアおよび核局在の組み合わせで、GFPを産生する。二つの顕著なコンパートメントはこれらの細胞中で容易に見分けることができ、元のGFP構造体のいずれかまたはどちらでもないの両者を発現する細胞の間の区別を容易にしている。
図1は、C.elegansにおいてRNA介在遺伝子阻害を研究するために使用された遺伝子を示す。RNA介在阻害を試験するために使用した遺伝子に対するイントロン・エクソン構造を示す(エクソン:塗りつぶしたます目;イントロン:空白のます目;5′および3′非翻訳領域:斜線;unc−229、unc−5412、fem−114、およびhIh−115)。これらの遺伝子は、(1)決定された分子構造、(2)ヌル表現型の性質を示す古典的遺伝子データ、を基準として選択された。阻害効果について試験された各セグメントは、遺伝子名に単文字が続く形で表わした(例えば、unc22C)。ゲノムDNA由来のセグメントは、遺伝子の上に示し、cDNA由来のセグメントは遺伝子の下に示す。これら遺伝子の各々に対する注入した二本鎖RNAの結果を、表1に記載する。各遺伝子のコード領域からのdsRNA配列は、その遺伝子に対するヌル表現型に類似した表現型を産生した。
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各RNAは6−10成体雌雄同体に注入された(0.5−1×106モルを各生殖腺腕部に)。4−6時間後(子宮から清澄な授精前の卵に)注入した動物を移し、20−22時間に卵を採取した。子孫表現型は孵化およびそれに続く12−24時間間隔でスコア化した。
a:RNA投与量と表現型応答の間の関係について半定量的な評価を得るため、我々は、一連の異なった濃度で各々のunc22ARNA調製物を注入した。試験した最高濃度(性腺当たり3.6×106モル)で、個々のセンスおよびアンチセンスunc22A調製物は幾分か識別できる痙攣(それぞれ子孫の1%および11%)を生じた。ds−unc22ARNAの比較投与で、全ての子孫で識別できる痙攣を生じたが、ds−unc22ARNAの120倍低い投与量で子孫の30%に識別できる痙攣を生じた。
b:unc22Cは、また、介在イントロン(43nt)を有する。
c:fem1Aもまた、イントロン10の部分(131nt)を有する。
d:最初の影響を受けた時期別産子群(注入後4−24時間に産卵)の動物は、unc−54のヌル突然変異体と区別できない運動欠損を示した。これらの動物のいくつかは(25−75%)産卵できなかった(unc−54ヌル突然変異体の別の表現型)が、しかし抑制動物の残りは産卵陽性であった。このことは、外陰筋におけるunc−54活性の部分的阻害を示している。後期の時期別産子群の動物は、しばしば体壁筋のサブセットに収縮性を伴った、著しい部分的機能喪失表現型を示す。
e:hlh−1阻害RNAの表現型は、ds−hlh1A注入体の事実上全ての子孫、およびds−hlh1Bとds−hlh1Cからの影響を受けた動物の約半数、にみられる休止胚および部分的に伸長したL1幼生(hlh−1ヌル表現型)ならびに(ds−hlh1Bおよびds−hlh1Cからの動物の残りにみられる)余り激しくない一連の欠損を含む。余り激しくない表現型はhlh−1の機能の部分的喪失の特徴である。
f:これらの注入宿主、PD4251、はミトコンドリアGFPおよび核GFP−LacZの両者を発現する。このことは、gfp(全ての蛍光を喪失)およびlacZ(核の蛍光を喪失)の阻害を同時に測定することが可能となる。表に、L1幼生としての動物のスコアリングを記載する。ds−gfpGは、これら幼生において、85体筋肉の0−3を除く全てでGFPの喪失をもたらした。これらの動物は成体に成熟するにつれて、GFP活性は0−5の別の体壁筋および8つの外陰筋においてみられた。
Claims (18)
- 哺乳類細胞内の標的遺伝子の発現を阻害する方法であって、リボ核酸(RNA)の、該標的遺伝子の発現を阻害するのに充分な量での該哺乳類細胞への導入を含み、該RNAは、該標的遺伝子のヌクレオチド配列にハイブリダイズすることができるリボヌクレオチド配列を含む第一の鎖と該第一の鎖に相補的なリボヌクレオチド配列を含む分離した第二の鎖との二本鎖構造であり、該第一および該第二の鎖配列は、互いにハイブリダイズして該二本鎖構造を形成する分離した相補鎖であり、そして該二本鎖構造は該標的遺伝子の発現を阻害する、
但し、細胞は、ヒト由来ではない、
ことを含む、方法。 - 標的遺伝子が内部遺伝子である請求項1記載の方法。
- 標的遺伝子が導入遺伝子である請求項1記載の方法。
- 標的遺伝子がウイルス遺伝子である請求項1記載の方法。
- 標的遺伝子のヌクレオチド配列にハイブリダイズすることができるリボヌクレオチド配列が少なくとも50塩基長である請求項1〜4のいずれか一項に記載の方法。
- 標的遺伝子発現が少なくとも10%阻害される請求項1〜5のいずれか一項に記載の方法。
- 哺乳類細胞が生物体内に存在し、標的遺伝子発現の阻害が機能表現型の喪失を表わす、請求項1〜6のいずれか一項に記載の方法。
- 標的遺伝子のヌクレオチド配列にハイブリダイズすることができるリボ核酸配列が少なくとも100塩基長である請求項1〜7のいずれか一項に記載の方法。
- 哺乳類細胞外での二本の相補鎖の合成およびRNA二本鎖形成の開始を更に含む請求項1〜7のいずれか一項に記載の方法。
- 哺乳類細胞内での二本の相補鎖の合成およびRNA二本鎖形成の開始を更に含む請求項1〜7のいずれか一項に記載の方法。
- 哺乳類細胞が生物体内に存在し、そしてRNAが生物体へ細胞外注入により導入される、請求項1〜9のいずれか一項に記載の方法。
- 哺乳類細胞内の発現構造体がRNAを産生する請求項1記載の方法。
- 標的遺伝子の発現を阻害する方法であって、
(a)標的哺乳類細胞を含有する生物体を提供することであって、該標的哺乳類細胞は該標的遺伝子を含有し、そして該標的遺伝子は該標的哺乳類細胞において発現される、そして
(b)該哺乳類細胞にリボ核酸(RNA)を提供することであって、該RNAは、該標的遺伝子のヌクレオチド配列にハイブリダイズすることができるリボヌクレオチド配列を含む第一の鎖と該第一の鎖に相補的なリボヌクレオチド配列を含む分離した第二の鎖との二本鎖構造であり、該第一および該第二の鎖配列は、互いにハイブリダイズして該二本鎖構造を形成する分離した相補鎖であり、そして該二本鎖構造は該哺乳類細胞内で該標的遺伝子の発現を阻害する、
但し、該生物体はヒトではない、
ことを含む、方法。 - 標的遺伝子のヌクレオチド配列に相補的なリボヌクレオチド配列が少なくとも100塩基長である請求項13記載の方法。
- 標的遺伝子のヌクレオチド配列に相補的なリボヌクレオチド配列が少なくとも50ヌクレオチド長である請求項13または14に記載の方法。
- 標的遺伝子発現が少なくとも10%阻害される請求項13〜15のいずれか一項に記載の方法。
- RNAが細胞外注入により生物体に導入される請求項13〜16のいずれか一項に記載の方法。
- 生物体内の遺伝子機能を同定する方法であって、
リボ核酸(RNA)の、標的哺乳類遺伝子の発現を阻害するのに充分な量での哺乳類細胞への導入を含み、該RNAは、該標的遺伝子のヌクレオチド配列にハイブリダイズすることができるリボヌクレオチド配列を含む第一の鎖と該第一の鎖に相補的なリボヌクレオチド配列を含む分離した第二の鎖との二本鎖構造であり、該第一および該第二の鎖配列は、互いにハイブリダイズして該二本鎖構造を形成する分離した相補鎖であり、そして該二本鎖構造は該標的遺伝子の発現を阻害する、ことを含む、方法によって、
該細胞内の該標的遺伝子の発現を阻害する、
但し、該細胞はヒト由来ではない、
ことを含む、方法。
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JP2002516062A (ja) | 2002-06-04 |
CA2311999A1 (en) | 1999-07-01 |
US8283329B2 (en) | 2012-10-09 |
US20030055020A1 (en) | 2003-03-20 |
US20160208280A1 (en) | 2016-07-21 |
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JP2015133993A (ja) | 2015-07-27 |
CA2311999C (en) | 2016-02-23 |
EP2267131B1 (en) | 2017-04-19 |
US20130029425A1 (en) | 2013-01-31 |
AU743798B2 (en) | 2002-02-07 |
JP5148028B2 (ja) | 2013-02-20 |
US6506559B1 (en) | 2003-01-14 |
EP2267131A1 (en) | 2010-12-29 |
US7560438B2 (en) | 2009-07-14 |
US20080081373A1 (en) | 2008-04-03 |
US10358653B2 (en) | 2019-07-23 |
US20130230492A1 (en) | 2013-09-05 |
US20030051263A1 (en) | 2003-03-13 |
US7538095B2 (en) | 2009-05-26 |
AU743798C (en) | 2006-02-09 |
EP1042462A1 (en) | 2000-10-11 |
US9102939B2 (en) | 2015-08-11 |
US20140350083A1 (en) | 2014-11-27 |
US8580754B2 (en) | 2013-11-12 |
JP2009291209A (ja) | 2009-12-17 |
JP2013048629A (ja) | 2013-03-14 |
US7622633B2 (en) | 2009-11-24 |
AU1938099A (en) | 1999-07-12 |
WO1999032619A1 (en) | 1999-07-01 |
US20080050342A1 (en) | 2008-02-28 |
US20030056235A1 (en) | 2003-03-20 |
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