JP7201153B2 - プログラム可能cas9-リコンビナーゼ融合タンパク質およびその使用 - Google Patents
プログラム可能cas9-リコンビナーゼ融合タンパク質およびその使用 Download PDFInfo
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Description
本出願は、35 U.S.C. § 119(e)の元で、2016年8月9日に出願された米国特許仮出願U.S.S.N. 62/372,755、および2017年2月7日に出願された米国特許仮出願U.S.S.N. 62/456,048の優先権を主張する;これらの各々は、参照により本明細書に組み込まれる。
本発明は、国立衛生研究所から授与された助成金番号R01EB022376およびR35GM118062の下、政府の支援を受けてなされた。政府は本発明において、一定の権利を有する。
効率的でプログラム可能および部位特異的な相同組換えは,遺伝学およびゲノム編集の長年の目標である。組換えを関心のある座位へ指向する初期の試みは、標的座位に相同な長い隣接配列を有するドナーDNAのトランスフェクションに頼っていた。この戦略は、非常に低い効率および、それ故に成分を同定するための厳格な選択の必要性により妨げられていた。より最近の努力は、二本鎖DNA切断(DSB)の、相同性指向修復(HDR)を誘導する能力を利用している。ホーミングエンドヌクレアーゼおよび後のプログラム可能なエンドヌクレアーゼ、例えばジンクフィンガーヌクレアーゼ、TALEヌクレアーゼ、Cas9、およびfCas9などは、標的化DSBを導入し、ドナーDNAの存在下でHDRを誘導するために使用されてきた。しかしながら、ほとんどの有糸分裂後細胞において、DSB誘導性のHDRは強く下方制御されており、一般に非効率的である。さらに、非相同末端結合(NHEJ)または一本鎖アニーリング(SSA)などの誤りがちな修復経路によるDSBの修復は、HDRよりも高い頻度で、DSB部位におけるヌクレオチドのランダムな挿入または欠失(インデル)を引き起こす。HDRの効率は、細胞が細胞周期の同期を強制する条件にさらされている場合、またはNHEJに関与する酵素が阻害されている場合に、高めることができる。しかし、かかる条件は、多くのランダムかつ予測不可能な事象を引き起こす可能性があり、潜在的な用途を制限する。本開示は、ガイドRNA特異的配列が隣接する最小リコンビナーゼコア部位を含むDNA部位を組換えることができ、内因性細胞機構または細胞状態からは独立した、未修飾細胞におけるプログラム可能な瘢痕のないゲノム編集へのステップを表す、融合タンパク質を提供する。
本開示は、ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメイン、任意のリンカー、およびリコンビナーゼ触媒ドメイン(例えば、Ginリコンビナーゼ触媒ドメインなどのセリンリコンビナーゼ触媒ドメイン、チロシンリコンビナーゼ触媒ドメイン、または任意の進化型リコンビナーゼ触媒ドメイン)を含む融合タンパク質の開発を記載する。この融合タンパク質は、2つのガイドRNA特異的DNA配列が隣接する最小gixコアリコンビナーゼ部位(NNNNAAASSWWSSTTTNNNN、配列番号19)で作動する。記載された融合タンパク質により媒介される組換えは、両方のガイドRNAに依存し、異なるガイドヌクレオチド:融合タンパク質複合体の間で直交性をもたらし、培養ヒト細胞において、ヒトゲノムで見出されるものと一致するDNA配列に対して効率的に機能する。本開示の融合タンパク質はまた、ヒト細胞(例えば、培養ヒト細胞)のゲノムに直接作用することができ、約14キロ塩基離れて位置する2つのrecCas9疑似部位間の欠失、挿入、反転、転座、または組換えを触媒する。この研究は、修飾されていないゲノムにおいてユーザ定義の単一塩基対分解を用いて、遺伝子挿入、欠失、反転、または染色体転座を触媒することができる、操作された酵素を提供する。
本明細書において単数形「a」、「an」、および「the」は、文脈から明らかにそうでないと示されない限り、単数形および複数形の言及を含む。したがって、例えば、「作用物質」への言及は、1つの作用物質および複数のかかる作用物質を含む。
dCas9(D10AおよびH840A):
例示の触媒的に不活性なCas9(dCas9):
CasX(uniprot.org/uniprot/F0NN87; uniprot.org/uniprot/F0NH53)
>tr|F0NN87|F0NN87_SULIH CRISPR関連Casxタンパク質OS=Sulfolobus islandicus(株HVE10/4)GN=SiH_0402 PE=4 SV=1
> APG80656.1CRISPR関連タンパク質CasY[未培養Parcubacteria群細菌]
本発明のこれらおよび他の態様の機能および利点は、以下の実施例からさらに十分に理解されるであろう。以下の実施例は、本発明の利点を説明することおよび特定の態様を説明することを意図しているが、本発明の全範囲を例示することは意図していない。したがって、実施例は本発明の範囲を限定することを意味しないことが理解されるであろう。
本明細書に記載の融合タンパク質および方法は、任意のプログラム可能DNA結合ドメインを使用することができる。
Cas9とDNA骨格間の静電相互作用が減少したCas9バリアント
野生型Francisella novicidaCpf1(配列番号714)(D917、E1006、およびD1255は太字かつ下線)
C2c1(uniprot.org/uniprot/T0D7A2#)
sp|T0D7A2|C2C1_ALIAG CRISPR関連エンドヌクレアーゼC2c1 OS=Alicyclobacillus acidoterrestris(株ATCC 49025/DSM 3922/CIP 106132/NCIMB 13137/GD3B)GN=c2c1 PE=1 SV=1
>sp|P0DOC6|C2C2_LEPSD CRISPR関連エンドリボヌクレアーゼC2c2 OS=Leptotrichia shahii(株DSM 19757/CCUG 47503/CIP 107916/JCM 16776/LB37)GN=c2c2 PE=1 SV=1
野生型Natronobacterium gregoryiアルゴノート(配列番号718)
本開示の組成物および方法において使用するためのリコンビナーゼ触媒ドメインは、任意のリコンビナーゼ由来であってよい。開示された方法および組成物における使用のための適切なリコンビナーゼ触媒ドメインは、例えば、限定はされないが、チロシンリコンビナーゼおよびセリンリコンビナーゼから得ることができる。本明細書で提供されるいくつかの例示の適切なリコンビナーゼとしては、例えば、限定はされないが、Ginリコンビナーゼ(gix部位に作用する)、Hinリコンビナーゼ(hix部位に作用する)、βリコンビナーゼ(six部位に作用する)、Sinリコンビナーゼ(resH部位に作用する)、Tn3リコンビナーゼ(res部位に作用する)、γδリコンビナーゼ(res部位に作用する)、バクテリオファージP1由来のCreリコンビナーゼ(LoxP部位に作用する);真菌由来のFLPリコンビナーゼ(FTR部位に作用する)。およびphiC31インテグラーゼ(att部位に作用する)が挙げられる。例示の適切なリコンビナーゼの非限定的な配列は、以下に見出され得る。
Creリコンビナーゼ配列
dCas9-36C6(ヌクレオチド)(配列番号765)
本開示の融合タンパク質は、本明細書に記載される要素の任意の組み合わせおよび順序であり得る。例示的な融合タンパク質としては、以下の構造のいずれかが挙げられるが、これらに限定はされない:NH2-[リコンビナーゼ触媒ドメイン]-[リンカー配列]-[ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメイン]-[任意のリンカー配列]-[任意のNLSドメイン]-[任意のリンカー配列]-[任意のアフィニティタグ]-COOH。別の態様において、融合タンパク質は次の構造を有する:NH2-[リコンビナーゼ触媒ドメイン]-[リンカー配列]-[ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメイン]-[任意のリンカー配列]-[NLSドメイン]-[任意リンカー配列]-[任意のアフィニティタグ]-COOH。別の態様において、融合タンパク質は次の構造を有する:NH2-[リコンビナーゼ触媒ドメイン]-[リンカー配列]-[ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメイン]-[任意のリンカー配列]-[NLSドメイン]-[任意のリンカー配列]-[アフィニティタグ]-COOH。別の態様において、融合タンパク質は次の構造を有する:NH2-[リコンビナーゼ触媒ドメイン]-[リンカー配列]-[ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメイン]-[リンカー配列]-[NLSドメイン]-[リンカー配列]-[アフィニティタグ]-COOH。
材料および方法
オリゴヌクレオチドおよびPCR
全てのオリゴヌクレオチドは、Integrated DNA Technologies(IDT, Coralville, CA)から購入したものであり、表1~5に列挙されている。酵素は、特記しない限り、New England Biolabs(Ipswich, MA)から購入した。プラスミドセーフATP依存性DNAseは、Epicenter(Madison, WI)から購入した。全てのアセンブルされたベクターを、One Shot Mach1-T1ファージ耐性化学的コンピテント細胞(Fisher Scientific, Waltham, MA)に形質転換した。特に明記しない限り、すべてのPCR反応はQ5 Hot Start High-Fidelity 2Xマスターミックスを用いて行った。Phusionポリメラーゼを、環状ポリメラーゼ伸長クローニング(CPEC)アセンブリに使用した。
表1.gRNA構築のためのオリゴヌクレオチド
5部構成のゴールデンゲートアセンブリを使用して、下記のレポーターを構築した。断片1~5はEsp3I部位に隣接していた;Esp3I消化は、断片の集合の順序を特定する相補的5’オーバーハングを作り出した(図6)。断片1、2、4、および5は、表5に示すフォワードおよびリバースの相補的オリゴヌクレオチドをアニーリングすることにより作製した。断片は、10μlの各オリゴヌクレオチド(100μM)を、20μlの分子等級水の中で混合し、95℃で3分間インキュベートし、温度を-0.1℃/秒の速度で16℃に下げることによってアニーリングした。断片3は、kanRおよびポリA終止コドンを含む領域を、プライマー3-forおよび3-revを用いてPCR増幅することにより作製した。これらのプライマーにはまた、Esp3Iが、この配列の5’および3’末端に付加された。
1)40~50ngの断片3
2)100ngのpCALNL EGFP-Esp3I
3)1μLのTango緩衝液(10倍)
4)1μLのDTT(10mM)
5)1μLのATP(10mM)
6)0.25μLのT7リガーゼ(3,000U/μL)
7)0.75μLのEsp3I(10U/μL)
8)10μLまでのH2O
特記しない限り、DNA断片は、アガロースゲルからQIAquickゲル抽出キット(Qiagen、Valencia, CA)を用いて単離し、DNA Clean & Concentrator-5(Zymo Research, Irvine, CA)またはQiaquick PCR精製キット(Qiagen, Valencia, CA)を用いてさらに精製した。ゲル精製を必要としないPCR断片を、上記のキットの1つを用いて単離した。
表8.gRNA配列のリスト
HEK293T細胞は、American Type Culture Collection(ATCC, Manassas, VA)から購入した。細胞を、10%ウシ胎児血清(FBS, Life Technologies, Carlsbad, CA)を添加したダルベッコの改変イーグル培地(DMEM)+GlutaMAX-1(4.5g/LのDグルコース+110mg/mLのピルビン酸ナトリウム)中で培養した。細胞を、37℃、5%CO2で加湿インキュベーター内で培養した。
トランスフェクションの60~72時間後、細胞をリン酸緩衝生理食塩水で洗浄し、50μLの0.05%トリプシン-EDTA(Life Technologies, Carlsbad, CA)で37℃で5~10分間回収した。細胞を250μLの培養培地で希釈し、BD Fortessaアナライザーで泳動した。iRFP蛍光を635nmのレーザーを用いて励起し、発光を670/30バンドパスフィルターを用いて収集した。488nMのレーザーを使用してEGFPを励起し、発光蛍光を505ロングパスフィルターおよび530/30バンドパスフィルターで取得した。データを、ライブイベントおよびトランスフェクトイベント(iRFPを発現する)についてゲートされたFlowJoソフトウェアで分析した。陽性GFP発現細胞を、少なくとも6,000のライブイベントからゲートされたトランスフェクト細胞の割合として測定した。最適化実験のために、アッセイバックグラウンドの決定を、recCas9またはガイドRNA発現ベクターなしで、レポータープラスミドとpUCとの同時トランスフェクションの際にeGFPを産生するトランスフェクト細胞のパーセンテージを測定することにより、行った。次いでこのバックグラウンドを、レポータープラスミドをrecCas9および標的または非標的ガイドRNA発現ベクターで同時トランスフェクトした場合に観察されたeGFP陽性細胞の割合から、差し引いた。
適切な標的部位の検索は、R統計プログラミングを用いたオープンソースバイオインフォマティクスパッケージであるBioconductor(Fu et al., High-frequency off-target mutagenesis induced by CRISPR-Cas nucleases in human cells. Nature biotechnology 31, 822-826 (2013);その全内容は参照により本明細書に組み込まれる)を用いて行った。Genome Reference Consortiumにより公開されたヒト参照ゲノムの最新リリース(GRCh38)を使用して、Cas9のPAM要件と本文に記載されている進化型gix配列の両方に一致する部位を検索した。ゲノムにRをロードすると、各検索パターンはBiostringとして表現された;これは、文字列のマッチングと操作を可能にするR内のコンテナーである。指定されたパラメータを使用して両方のDNA鎖を全ゲノムについてスキャンすると、約450の潜在的標的が、GRCh38参照アセンブリを使用して検索した場合のヒトゲノムにおいて明らかになった(表9)。
表9.in silicoで同定されたrecCas9ゲノム標的
293T細胞のトランスフェクションを上記のように6重で実施し、72時間インキュベートした。細胞を採取し、複製物を合わせた。エピソームDNAを、アルカリ溶解およびスピンカラム精製を含む修正HIRT抽出を用いて、本質的に記載されているようにして抽出した(Quan et al., Circular polymerase extension cloning of complex gene libraries and pathways. PloS one 4, e6441 (2009);およびHillson (2010), vol. 2015, pp. CPEC protocol;これら各々の全内容は参照により本明細書に組み込まれる)。簡単に説明すると、採取後、HEK293T細胞を500μLの氷冷PBSで洗浄し、250μLのGTE緩衝液(50mMグルコース、25mMのTris-HCl、10mMのEDTAおよびpH8.0)に再懸濁し、室温で5分間インキュベートし、氷上で5分間、200μLの溶解緩衝液(200mMのNaOH、1%ドデシル硫酸ナトリウム)を用いて溶解した。溶解を150μLの酢酸カリウム溶液(5M酢酸塩、3Mカリウム、pH6.7)で中和した。細胞破片を、21,130gで15分間の遠心分離によりペレット化し、溶解物をEconospin Spinカラム(Epoch Life Science, Missouri City, TX)に供した。カラムを750μLの洗浄緩衝液(Omega Bio-tek, Norcross, GA)で2回洗浄し、45μLのTE緩衝液、pH8.0中で溶出した。
HEK293T細胞を、1ウェルあたり6×105細胞の密度で24ウェルコラーゲン処理プレートに播種し、一晩増殖させた(Corning, Corning, NY)。トランスフェクション反応は、Opti-MEM(ThermoFisher Scientific, Waltham, MA)中で最終容量100μLにした。各トランスフェクションについて、90ngの各ガイドRNA発現ベクター、20ngのpmaxGFP(Lonza, Allendale, NJ)および320ngのrecCas9発現ベクターを、Opti-MEM(ThermoFisher Scientific, Waltham, MA)中の2μLのリポフェクタミン2000と混合し、個々のウェルに加えた。48時間後、細胞を採取し、GFPトランスフェクション対照について、BD FACS AriIIIuセルソーター上で選別した。細胞を純度モードで100μmのノズルを用いて選別し、バックグラウンドの蛍光を、トランスフェクトしていない細胞との比較により決定した。選別した細胞をPBS中の氷上で収集し、ペレット化し、冷PBSで2回洗浄した。ゲノムDNAは、E.Z.N.A. Tissue DNAキット(Omega Bio-Tek, Norcross, GA)を用いて採取し、100μLのEBで溶出した。ゲノムDNAを、Tecan Infinite M1000 Pro蛍光プレートリーダー上で測定するQuant-iT PicoGreen dsDNAキット(ThermoFisher Scientific, Waltham, MA)を用いて定量した。
GinリコンビナーゼをdCas9に融合する
最近、dCas9のN末端がFokIヌクレアーゼ触媒ドメインに融合すると、2つのガイドRNA特異的配列に隣接するDNA部位を切断する二量体dCas9-FokI融合物をもたらし得ることが実証された(例えば、下記参照:Guilinger et al., Fusion of catalytically inactive Cas9 to FokI nuclease improves the specificity of genome modification. Nature biotechnology, (2014);Tsai et al., Dimeric CRISPR RNA-guided FokI nucleases for highly specific genome editing. Nature biotechnology (2014);これら各々の全内容は、参照により本明細書に組み込まれる)。同じ融合配向を用いて、dCas9を、以前にBarbasらによって進化された二量体Ginインベルターゼの高活性触媒ドメインであるGinβに結合させた(Gaj et al., A comprehensive approach to zinc-finger recombinase customization enables genomic targeting in human cells. Nucleic acids research 41, 3937-3946 (2013);その全内容は参照により本明細書に組み込まれる)。Ginβは、天然コア配列CTGTAAACCGAGGTTTTGGA(配列番号700)に関連するいくつかの20bpコア「gix」配列を、無差別に組換える(Gaj et al., A comprehensive approach to zinc-finger recombinase customization enables genomic targeting in human cells. Nucleic acids research 41, 3937-3946 (2013);Klippel et al., The DNA Invertase Gin of Phage Mu - Formation of a Covalent Complex with DNA Via a Phosphoserine at Amino-Acid Position-9. Embo Journal 7, 1229-1237 (1988);Mertens et al., Site-specific recombination in bacteriophage Mu: characterization of binding sites for the DNA invertase Gin. The EMBO journal 7, 1219-1227 (1988);Plasterk et al., DNA inversions in the chromosome of Escherichia coli and in bacteriophage Mu: relationship to other site-specific recombination systems. Proceedings of the National Academy of Sciences of the United States of America 80, 5355-5358 (1983);これら各々の全内容は参照により本明細書に組み込まれる)。ガイドRNAは、recCas9二量体を2つのガイドRNA特異的配列が隣接するgix部位に局在化し、DNA組換えを、GinβドメインがガイドRNAプログラム化様式で触媒することを可能にする(図1D)。
観察された低レベルの活性は、最適以下のガイドRNA配列またはコアgix配列によって引き起こされ得、これは、ガイドRNA:Cas9結合の効率が配列依存的であることを示す以前の報告と一致する(例えば、Xu et al., Sequence determinants of improved CRISPR sgRNA design. Genome research 25, 1147-1157 (2015)を参照、この全内容は参照により本明細書に組み込まれる)。さらに、本最適化は天然のgixコア配列を用いて実施されたが(例えば、Klippel et al., The DNA Invertase Gin of Phage Mu - Formation of a Covalent Complex with DNA Via a Phosphoserine at Amino-Acid Position-9. Embo Journal 7, 1229-1237 (1988);Mertens et al., Site-specific recombination in bacteriophage Mu: characterization of binding sites for the DNA invertase Gin. The EMBO journal 7, 1219-1227 (1988);Plasterk et al., DNA inversions in the chromosome of Escherichia coli and in bacteriophage Mu: relationship to other site-specific recombination systems. Proceedings of the National Academy of Sciences of the United States of America 80, 5355-5358 (1983)を参照;これら各々の全内容は参照により本明細書に組み込まれる)、しかしいくつかの研究は、ジンクフィンガー-Gin融合体またはTALE-Gin融合体が活性であり、ある場合にはわずかに変更されたコア部位において、より活性であることを示した。例えば、以下を参照:Gordley et al., 3rd, Synthesis of programmable integrases. Proceedings of the National Academy of Sciences of the United States of America 106, 5053-5058 (2009);Gersbach et al., Targeted plasmid integration into the human genome by an engineered zinc-finger recombinase. Nucleic acids research 39, 7868-7878 (2011);Mercer et al., Chimeric TALE recombinases with programmable DNA sequence specificity. Nucleic acids research 40, 11163-11172 (2012);Gaj et al., A comprehensive approach to zinc-finger recombinase customization enables genomic targeting in human cells. Nucleic acids research 41, 3937-3946 (2013);Gordley et al., 3rd, Evolution of programmable zinc finger-recombinases with activity in human cells. J Mol Biol 367, 802-813 (2007);Gersbach et al., 3rd, Directed evolution of recombinase specificity by split gene reassembly. Nucleic acids research 38, 4198-4206 (2010);およびGaj et al., Structure-guided reprogramming of serine recombinase DNA sequence specificity. Proceedings of the National Academy of Sciences of the United States of America 108, 498-503 (2011);これら各々の全内容は参照により本明細書に組み込まれる。したがって、未修飾のヒトゲノム配列がrecCas9によって標的化され得るかどうかを試験するため、およびガイドRNAおよびコア配列を変化させることがrecCas9活性を増加させるかどうかを試験するために、ヒトゲノム内に見出される配列を標的とした。
次に、recCas9が、ヒトゲノム中に見出される複数の別々の座位に一致する配列を直交する様式で標的とし得るかどうかを試験した。ヒトゲノム中のrecCas9標的部位のサブセットの選択を、ゲノム組込みのためのセーフハーバー座位としてのそれらの潜在的使用に基づき、またはある場合には遺伝病に関与する遺伝子内のそれらの位置に基づいて、行った。
レポータープラスミドのrecCas9媒介組換えの産物を特徴付けして、EGFP発現がポリAターミネーター配列のrecCas9媒介性除去の結果であることを確認した。レポータープラスミドを、recCas9発現ベクターと同族または非同族ガイドRNA対を産生するプラスミドとの同時トランスフェクションの後に、5番染色体部位1、12番染色体、および13番染色体(FGF14座位)について配列決定した。72時間のインキュベーション後、エピソームDNAを抽出し(上記の通り)、大腸菌に形質転換してレポータープラスミドを単離した。レポータープラスミドを含有する単一コロニーを配列決定した(図4B)。
最後に、recCas9が、培養ヒト細胞のゲノムDNAに対して直接作用することができるかどうかを調べた。ヒトゲノム中の潜在的なrecCas9認識部位のリスト(表9)を使用して、recCas9によって標的化された場合にPCRにより検出可能な染色体欠失事象を生じるであろう部位の対を探した。ガイドRNA発現ベクターは、5番染色体部位1または13番染色体(FGF14座位)、すなわち両者とも一過性トランスフェクションアッセイにおいて組み換えられることが示された部位に、recCas9を導くように設計された(図4)。新しい標的部位は、5番染色体部位1の約3から23Mbp上流および7から10Mbp下流、および13番染色体FGF14部位の12から44Mbp上流の範囲であった。recCas9発現ベクターを、これらの新しいガイドRNA対および5番染色体部位1または13番染色体FGF14に使用される検証済みガイドRNA対のそれぞれと、同時トランスフェクトしたが、ゲノムPCRによる染色体欠失の証拠は観察されなかった。
「36C6」と呼ばれるヒトゲノムのRosa座位内の部位を標的とするように進化させたCreリコンビナーゼを、dCas9に融合した。次いで、この融合物を使用して、Rosa標的部位を含有するプラスミドベースのレポーターを、ガイドRNA依存性様式で組み換えた。図7Aは、野生型Creおよび36C6を用いたリンカー最適化の結果を示す。示されている1×2×、5×、および8×リンカーは、リンカー中のGGS反復の数である。復帰変異分析(reversion analysis)は、dCas9に融合した36C6に変異を作ることが、キメラ融合物の相対的ガイド依存性に影響を及ぼし得ることを実証した(図7B)。復帰変異は、それらの非変異アミノ酸で標識されている。例えば、Mに変異した306位は、アッセイが行われる前にIに戻された。同族レポーターを標的とするGinB構築物を、図7Aおよび7Bに示される実験データのための対照として使用した。オンターゲットのガイドは、chr13-102010574ガイド(プラスミドBC165および166)であった。示されている略語は、GGS-36C6:dCas9-GGS-36C6;2GGS-36C6(リンカー配列番号182を使用):sdCas9-GGSGGS-36C6(リンカー配列番号182を使用)である。
参考文献
Claims (28)
- 以下:
(i) ガイドRNA(gRNA)に結合しているヌクレアーゼ不活性Cas9(dCas9)ドメイン;
(ii) (GGS)8(配列番号183)を含むリンカー;
(iii) 配列番号713と少なくとも95%の配列同一性を有するアミノ酸配列を有し、かつ配列番号713の127Lに対応するアミノ酸を含み、任意に、配列番号713のアミノ酸106Y、136R、および137Fに対応する少なくとも1つのアミノ酸を含む、gixコアまたはgix関連コアの配列に結合するGinリコンビナーゼ触媒ドメイン;ならびに
(iv) 核局在化シグナル(NLS)ドメインを含む、融合タンパク質であって、
前記融合タンパク質が、構造:NH2-[(iii)Ginリコンビナーゼ触媒ドメイン]-[(ii)リンカー配列]-[(i)dCas9ドメイン]-[任意のリンカー配列]-[(iv)NLSドメイン]-COOHを含む;
dCas9ドメインのアミノ酸配列が、配列番号1のD10Aおよび/またはH840A変異に対応する変異を含むか、あるいはdCas9ドメインのアミノ酸配列が、配列番号712と95%以上の配列同一性を有する;ならびに
gRNAが、gixコアまたはgix関連コアの配列から3~6ヌクレオチド離れた標的配列に結合する、前記融合タンパク質。 - dCas9ドメインのアミノ酸配列が、配列番号1に示されるN末端メチオニンを含まない、請求項1に記載の融合タンパク質。
- dCas9ドメインが、配列番号712である、請求項1または2に記載の融合タンパク質。
- Ginリコンビナーゼ触媒ドメインのアミノ酸配列が、配列番号713のアミノ酸配列を含む、請求項1~3のいずれか一項に記載の融合タンパク質。
- NLSドメインが、dCas9ドメインに、第2のリンカーを介して結合している、請求項1~4のいずれか一項に記載の融合タンパク質。
- 第2のリンカーが、
(a) ペプチドリンカー;
(b) アミノ酸配列SGSETPGTSESATPES(配列番号7)、SGSETPGTSESA(配列番号8)、もしくはSGSETPGTSESATPEGGSGGS(配列番号9)を有するXTENリンカー;
(c) トリペプチドGGSの1個以上の反復(任意に、6~10個の反復もしくは8個の反復を包含する)を含むアミノ酸配列;
(d) アミノ酸配列GGSGGSGGSGGSGGSGGSGGSGGS(配列番号183);
(e) VPFLLEPDNINGKTC(配列番号10)、GSAGSAAGSGEF(配列番号11)、SIVAQLSRPDPA(配列番号12)、MKIIEQLPSA(配列番号13)、VRHKLKRVGS(配列番号14)、GHGTGSTGSGSS(配列番号15)、MSRPDPA(配列番号16)、もしくはGGSM(配列番号17)を含むアミノ酸配列;または
(f) ポリエチレングリコール(PEG)、ポリプロピレングリコール(PPG)、コポリ(エチレン/プロピレン)グリコール、ポリオキシエチレン(POE)、ポリウレタン、ポリホスファゼン、多糖類、デキストラン、ポリビニルアルコール、ポリビニルピロリドン、ポリビニルエチルエーテル、ポリアクリルアミド、ポリアクリレート、ポリシアノアクリレート、脂質ポリマー、キチン、ヒアルロン酸、ヘパリン、もしくはアルキルリンカーを含む、非ペプチドリンカーである、請求項5に記載の融合タンパク質。 - 配列番号719もしくは配列番号185に示されるアミノ酸配列か、または配列番号719もしくは配列番号185と95%以上の配列同一性を有するアミノ配列を含む、請求項1~6のいずれか一項に記載の融合タンパク質。
- 1以上のアフィニティタグをさらに含み、任意に、1以上のアフィニティタグが、FLAGタグ、ポリヒスチジン(ポリHis)タグ、ポリアルギニン(ポリArg)タグ、Mycタグ、およびHAタグからなる群から選択される、請求項1~7のいずれか一項に記載の融合タンパク質。
- 請求項1~8のいずれか一項に記載の融合タンパク質の、二量体。
- DNA分子に結合している、請求項9に記載の二量体。
- 二量体の各融合タンパク質が、DNA分子の同じ鎖に結合している、請求項10に記載の二量体。
- 二量体の各融合タンパク質が、DNA分子の対向鎖に結合している、請求項10に記載の二量体。
- 二量体のgRNAが、リコンビナーゼ部位に隣接する標的配列にハイブリダイズする、請求項9~12のいずれか一項に記載の二量体。
- リコンビナーゼ部位が、gixコアまたはgix関連コアの配列を含む、請求項13に記載の二量体。
- gixコアまたはgix関連コアの配列と少なくとも1つの標的配列との間の距離が、5~6塩基対である、請求項13に記載の二量体。
- 第1の二量体が第2の二量体に結合し、それによって融合タンパク質の四量体を形成する、請求項9~15のいずれか一項に記載の二量体。
- 請求項1~8のいずれか一項に記載の融合タンパク質の四量体であって、任意に、四量体は、DNA分子に結合しており、および、四量体の各二量体は、DNAの対向鎖またはDNAの同じ鎖に結合している、前記四量体。
- 2つのDNA分子間の部位特異的組換え方法であって、
(a) 第1のDNAを第1の融合タンパク質と接触させること
(ここで、ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、第1のDNAの第1の標的配列にハイブリダイズする第1のgRNAに結合する);
(b) 第1のDNAを第2の融合タンパク質と接触させること
(ここで、第2の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、第1のDNAの第2の標的配列にハイブリダイズする第2のgRNAに結合する);
(c) 第2のDNAを第3の融合タンパク質と接触させること
(ここで、第3の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、第2のDNAの第1の標的配列にハイブリダイズする第3のgRNAに結合する);および
(d) 第2のDNAを第4の融合タンパク質と接触させること
(ここで、第4の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、第2のDNAの第2の標的配列にハイブリダイズする第4のgRNAに結合する)を含み、
ここで、ステップ(a)~(d)における融合タンパク質の結合は、DNAが組み換えされるような条件下で、融合タンパク質のリコンビナーゼ触媒ドメインの四量体化をもたらし、
ここで、前記DNAがヒト体内になく、ならびに
ここで、第1、第2、第3、および/または第4の融合タンパク質は、請求項1~8のいずれか一項に記載の融合タンパク質である、前記方法。 - 単一のDNA分子の2つの領域間の部位特異的組換え方法であって、
(a) DNAを第1の融合タンパク質と接触させること
(ここで、ガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、DNAの第1の標的配列にハイブリダイズする第1のgRNAに結合する);
(b) DNAを第2の融合タンパク質と接触させること
(ここで、第2の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、DNAの第2の標的配列にハイブリダイズする第2のgRNAに結合する);
(c) DNAを第3の融合タンパク質と接触させること
(ここで、第3の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインが、DNAの第3の標的配列にハイブリダイズする第3のgRNAに結合する);および
(d) DNAを第4の融合タンパク質と接触させること
(ここで、第4の融合タンパク質のガイドヌクレオチド配列プログラム可能DNA結合タンパク質ドメインは、DNAの第4の標的配列にハイブリダイズする第4のgRNAに結合する)を含み、
ここで、ステップ(a)~(d)における融合タンパク質の結合は、DNAが組み換えされるような条件下で、融合タンパク質のリコンビナーゼ触媒ドメインの四量体化をもたらし、
ここで、前記DNAがヒト体内になく、ならびに
ここで、第1、第2、第3、および/または第4の融合タンパク質は、請求項1~8のいずれか一項に記載の融合タンパク質である、前記方法。 - ステップ(a)および(b)のgRNA、および/またはステップ(c)および(d)のgRNAが、リコンビナーゼ部位に隣接する標的配列にハイブリダイズする、請求項19に記載の方法。
- リコンビナーゼ部位が、gixコアまたはgix関連コアの配列を含む、請求項20に記載の方法。
- gixコアまたはgix関連コアの配列と少なくとも1つのgRNA結合部位との間の距離が、5~6塩基対である、請求項21に記載の方法。
- 請求項1~8(請求項6(f)を除く)のいずれか一項に記載の融合タンパク質をコードし、および前記融合タンパク質のヌクレアーゼ不活性Cas9(dCas9)ドメインに結合する前記gRNAをコードする、少なくとも1つのポリヌクレオチド。
- 請求項23に記載のポリヌクレオチドを含む、ベクター。
- 請求項1~8(請求項6(f)を除く)のいずれか一項に記載の融合タンパク質をコードし、および前記融合タンパク質のヌクレアーゼ不活性Cas9(dCas9)ドメインに結合する前記gRNAをコードする少なくとも1つのポリヌクレオチドを含む、組換えタンパク質発現用ベクター。
- 請求項1~8(請求項6(f)を除く)のいずれか一項に記載の融合タンパク質と、前記融合タンパク質のヌクレアーゼ不活性Cas9(dCas9)ドメインに結合する前記gRNAとを発現するための遺伝子構築物を含む、細胞。
- gixコアまたはgix関連コアの配列が、ヌクレオチド配列NNNNAAASSWWSSTTTNNNN(配列番号19)を含み、ここで各Nは独立して任意のヌクレオチドであり、WはAまたはTであり、およびSはGまたはCである、請求項1~8のいずれか一項に記載の融合タンパク質。
- gixコアリコンビナーゼ部位が、ヌクレオチド配列CTGTAAACCGAGGTTTTGGA(配列番号700)を含む、請求項1~8のいずれか一項に記載の融合タンパク質。
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JP2015532654A (ja) | 2012-09-04 | 2015-11-12 | ザ スクリプス リサーチ インスティテュート | 標的結合特異性を有するキメラポリペプチド |
WO2015035162A2 (en) | 2013-09-06 | 2015-03-12 | President And Fellows Of Harvard College | Cas9 variants and uses thereof |
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JP2022122919A (ja) | 2022-08-23 |
US20240209329A1 (en) | 2024-06-27 |
CA3033327A1 (en) | 2018-02-15 |
JP2019526248A (ja) | 2019-09-19 |
AU2017308889B2 (en) | 2023-11-09 |
US11661590B2 (en) | 2023-05-30 |
EP3497214A1 (en) | 2019-06-19 |
EP3497214B1 (en) | 2023-06-28 |
AU2017308889A1 (en) | 2019-02-28 |
WO2018031683A1 (en) | 2018-02-15 |
US20190367891A1 (en) | 2019-12-05 |
CN109804066A (zh) | 2019-05-24 |
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