KR20210139271A - Crispr/cas 융합 단백질 및 시스템 - Google Patents
Crispr/cas 융합 단백질 및 시스템 Download PDFInfo
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- KR20210139271A KR20210139271A KR1020217029732A KR20217029732A KR20210139271A KR 20210139271 A KR20210139271 A KR 20210139271A KR 1020217029732 A KR1020217029732 A KR 1020217029732A KR 20217029732 A KR20217029732 A KR 20217029732A KR 20210139271 A KR20210139271 A KR 20210139271A
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Abstract
조작된 Cas9 시스템들이 본원에 개시되어 있다.
Description
관련 출원
본 출원은 2019년 2월 15일 출원된 미국 가특허출원 제 62/806,708호의 우선권의 이익을 주장하며, 이 문헌은 본 명세서에 그 전문이 참고문헌으로 포함된다.
기술분야
본 발명은 조작된 Cas9 시스템, 이러한 시스템을 인코딩하는 핵산, 및 게놈 변형을 위해 이러한 시스템을 사용하는 방법에 관한 것이다.
배경
GFP를 Cas9에 융합하기 위해 다양한 유형의 펩티드 링커가 테스트되었지만 통상적으로 기본 Cas9의 활성을 더 낮춘다.
발명의 요약
본 개시내용의 다양한 양상들 중에는 조작된 Cas9 시스템이 포함된다.
본 발명의 다른 양상들 및 특징들을 이하에서 상세히 설명한다.
도면의 간단한 설명
도 1은 본 명세서에 개시된 Cas9 융합 단백질이 각각 SpCas9 단백질 수준과 유사한 편집 활성을 유지함을 보여준다.
도 2A 및 도 2B는 본원에 공개된 Cas9 융합 단백질의 편집 효율이 모든 표적에서 시판 단백질의 편집 효율보다 몇 배 더 높았음을 보여준다.
서열 목록
본원은 ASCII 형식으로 전자적으로 제출된 서열 목록을 포함하며, 이러한 서열 목록은 본원에 그 전문이 참조로 포함된다. 2020년 2월 13일 생성된 상기 ASCII 사본 파일명은 P19-027_WO-PCT_SL.txt이며 87,735 바이트 크기이다.
도 1은 본 명세서에 개시된 Cas9 융합 단백질이 각각 SpCas9 단백질 수준과 유사한 편집 활성을 유지함을 보여준다.
도 2A 및 도 2B는 본원에 공개된 Cas9 융합 단백질의 편집 효율이 모든 표적에서 시판 단백질의 편집 효율보다 몇 배 더 높았음을 보여준다.
서열 목록
본원은 ASCII 형식으로 전자적으로 제출된 서열 목록을 포함하며, 이러한 서열 목록은 본원에 그 전문이 참조로 포함된다. 2020년 2월 13일 생성된 상기 ASCII 사본 파일명은 P19-027_WO-PCT_SL.txt이며 87,735 바이트 크기이다.
상세한 설명
CRISPR 단백질에 대한 부속 단백질의 융합은 다양한 단백질 기능을 세포 내의 정의된 위치에 국소화시킬 수 있는 광범위한 기회를 생성한다. 무엇보다도, CRISPR 기능을 보존하는 방식으로 CRISPR 단백질에 이종 단백질의 융합을 가능하게 하는 펩티드 링커가 개시되어 있다.
(I)
조작된 Cas9 시스템
본 발명의 한 양상은 조작된 Cas9 단백질 및 시스템을 제공한다. 예를 들어, Cas9-마커 융합 단백질이 개시되어 있다. 일부 양상에서, 시스템은 조작된 Cas9 단백질 및 조작된 가이드 RNA를 포함하며, 여기서 각각의 조작된 가이드 RNA는 특이적으로 조작된 Cas9 단백질과 복합체를 형성하도록 설계된다. 이러한 조작된 Cas9 시스템은 자연적으로 발생하지 않는다.
(a) 조작된 Cas9 단백질
Cas9 단백질은 다양한 박테리아에 존재하는 II형 CRISPR 시스템의 단일 이펙터 단백질이다. 본원에 개시된 조작된 Cas9 단백질은 아카리오클로리스 종 (Acaryochloris sp.), 아세토할로비움 종 (Acetohalobium sp.), 아시다미노코쿠스 종 (Acidaminococcus sp.), 아시디티오바실루스 종 (Acidithiobacillus sp.), 아시도테르무스 종 (Acidothermus sp.), 아커만시아 종 (Akkermansia sp.), 알리사이클로바실루스 종 (Alicyclobacillus sp.), 알로크로마티움 종 (Allochromatium sp.), 암모니펙스 종 (Ammonifex sp.), 아나바에나 종 (Anabaena sp.), 아르트로스피라 종 (Arthrospira sp.), 바실루스 종 (Bacillus sp.), 비피도박테릴움 종 (Bifidobacterium sp.), 부르크홀데리알레스 종 (Burkholderiales sp.), 칼디셀룰로시룹터 종 (Caldicelulosiruptor sp.), 캄필로박터 종 (Campylobacter sp.), 칸디다투스 종 (Candidatus sp.), 클로스트리디움 종 (Clostridium sp.), 코리네박테리움 종 (Corynebacterium sp.), 크로코스파에라 종 (Crocosphaera sp.), 시아노테세 종 (Cyanothece sp.), 엑시구오박테리움 종 (Exiguobacterium sp.), 피브로박터 종 (Fibrobacter sp.), 피네골디아 종 (Finegoldia sp.), 프란시셀라 종 (Francisella sp.), 크테도노박터 종 (Ktedonobacter sp.), 라크노스피라세아에 종 (Lachnospiraceae sp.), 락토바실루스 종 (Lactobacillus sp.), 리스테리아 종 (Listeria sp.), 린그비아 종 (Lyngbya sp.), 마리노박터 종 (Marinobacter sp.), 메타노할로비움 종 (Methanohalobium sp.), 미크로실라 종 (Microscilla sp.), 미크로콜레우스 종 (Microcoleus sp.), 미크로시스티스 종 (Microcystis sp.), 마이코플라즈마 종 (Mycoplasma sp.), 나트라나에로비우스 종 (Natranaerobius sp.), 네이세리아 종 (Neisseria sp.), 니트라티프락터 종 (Nitratifractor sp.), 니트로소코쿠스 종 (Nitrosococcus sp.), 노카르디옵시스 종 (Nocardiopsis sp.), 노둘라리아 종 (Nodularia sp.), 노스톡 종 (Nostoc sp.), 오에노코쿠스 종 (Oenococcus sp.), 오실라토리아 종 (Oscillatoria sp.), 파라수테렐라 종 (Parasutterella sp.), 파스퇴렐라 종 (Pasteurella sp.), 파르비바쿨룸 종 (Parvibaculum sp.), 펠로토마쿨룸 종 (Pelotomaculum sp.), 페트로로가 종 (Petrotoga sp.), 폴라로모나스 종 (Polaromonas sp.), 프레보텔라 종 (Prevotella sp.), 수도알테로모나스 종 (Pseudoalteromonas sp.), 랄스토니아 종 (Ralstonia sp.), 로도스피릴룸 종 (Rhodospirillum sp.), 스타필로코쿠스 종 (Staphylococcus sp.), 스트렙토코쿠스 종 (Streptococcus sp.), 스트렙토마이세스 종 (Streptomyces sp.), 스트렙토스포란지움 종 (Streptosporangium sp.), 시네코코쿠스 종 (Synechococcus sp.), 테르모시포 종 (Thermosipho sp.), 트레포네마 종 (Treponema sp.), 베루코미크로비아 종 (Verrucomicrobia sp)., 및 올리넬라 종 (Wolinella sp.)에서 유래할 수 있다.
Cas9 단백질 또는 다른 구성성분들이 유래 또는 유도될 수 있는 예시적인 종들에는 아카리오클로리스 종 (예를 들어, 아카리오클로리스 마리나), 아세토할로비움 종 (예컨대, 아세토할로비움 아라바티쿰), 아시다미노코쿠스 종, 아시디티오바실루스 종 (예를 들어, 아시디티오바실루스 칼두스, 아시디티오바실루스 페로옥시단스), 아시도테르무스 종, 아케르만시아 종, 알리사이클로바실루스 종 (예를 들어, 알리사이클로바실루스 아시도칼다리우스), 알로크로마티움 종 (예를 들어, 알로크로마티움 비노숨), 암모니펙스 종 (예를 들어, 암모니펙스 데겐시이), 아나바에나 종 (예를 들어, 아나바에나 바리아빌리스), 아르트로스피라 종 (예를 들어, 아르트로스피라 맥시마, 아르트로스피라 플라텐시스), 바실루스 종 (예를 들어, 바실루스 수도마이코이데스, 바실루스 셀레니티레두센스), 비피도박테리움 종, 벌크홀데리알레스 종 (예를 들어, 벌크홀데리알레스 박테리움), 칼디셀룰로시럽터 종 (예를 들어, 칼디셀룰로시럽터 벡시이), 캄필로박터 종 (예를 들어, 캄필로박터 제주니, 캄필로박터 라리), 칸디다투스 종, (예를 들어, 칸디다투스 데술포루디스), 클로스트리디움 종 (예를 들어, 클로스트리디움 보툴리눔, 클로스트리디움 디피실레), 코리네박테리움 종 (예를 들어, 코리네박테리움 디프테리아), 크로코스파에라 종 (예를 들어, 크로코스파에라 왓소니이), 시아노테세 종, 델타프로테오박테리움 종, 엑시구오박테리움 종 (예를 들어, 엑시구오박테리움 시비리쿰), (피브로박터 종 (예를 들어, 피브로박터 숙시노겐), 피네골디아 종 (예를 들어, 피네골디아 마그나), 프란시셀라 종 (예를 들어, 프란시셀라 노비시다), 감마프로테오박테리움, 크테도노박터 종 (예를 들어, 크테도노박터 라세미페르), 라크노스피라세아에 종, 락토바실루스 종 (예를 들어, 락토바실루스 부크네리, 락토바실루스 델브루엑키이, 락토바실루스 가세리, 락토바실루스 살리바리우스), 리스테리아 종 (예를 들어, 리스테리아 이노쿠아), 렙토트리키아 종, 린그비아 종, 마리노박터 종, 메타노할로비움 종 (예를 들어, 메타노할로비움 에베스티가툼), 마이크로콜레우스 종 (예를 들어, 마이크로콜레우스 크토노플라스테스), 마이크로실라 종 (예를 들어, 마이크로실라 마리나), 마이크로시스티스 종 (예를 들어, 마이크로시스티스 아에루기노사), 마이코플라스마 종, 나트라나에로비우스 종 (예를 들어, 나트라나에로비우스 써모필루스), 네이세리아 종 (예를 들어, 네이세리아 시네레아, 네이세리아 메닝기티디스), 니트라티프락터 종, 니트로소코쿠스 종 (예를 들어, 니트로소모무스 할로필루스, 니트로소코쿠스 왓소니), 노카르디옵시스 종 (예를 들어, 노카르디옵시스 다쏜빌레이), 노둘라리아 종 (예를 들어, 노둘라리아 스푸미게나), 노스톡 종, 오에노코쿠스 종, 오실라토리아 종, 파라수테렐라 종, 파비바쿨럼 종 (예를 들어, 파비바쿨럼 라바멘티보란스), 파스퇴렐라 종 (예를 들어, 파스퇴렐라 파스퇴렐라 멀토시다), 펠로토마쿨럼 종, (예를 들어, 펠로토마쿨럼 써모프로피오니쿰), 페트로토가 종 (예를 들어, 페트로토가 모빌리스), 플랑크토마이세스 종, 폴라로모나스 종 (예를 들어, 폴라로모나스 나프탈레니보란스), 프레보텔라 종, 수도알테로모나스 종 (예를 들어, 수도알테로모나스 할로플랑크티스), 랄스토니아 종, 루미노코쿠스 종, 로도스피릴룸 종 (예를 들어, 로도스피릴룸 루브룸), 스타필로코쿠스 종 (예를 들어, 스타필로코쿠스 아우레우스), 스트렙토코쿠스 종 (예를 들어, 스트렙토코쿠스 파스퇴리아누스, 스트렙토코쿠스 피오게네스, 스트렙토코쿠스 써모필루스), 수테렐라 종 (예를 들어, 수테렐라 와즈워텐시스), 스트렙토마이세스 종 (예를 들어, 스트렙토마이세스 프리스티나에스피랄리스, 스트렙토마이세스 비리도크로모게네스, 스트렙토마이세스 비리도크로모게네스), 스트렙토스포란기움 종 (예를 들어, 스트렙토스포란기움 로세움, 스트렙토스포란기움 로세움), 시네코코쿠스 종, 써모시포 종 (예를 들어, 써모시포 아프리카누스), 트레포네마 종 (예를 들어, 트레포네마 덴티콜라), 및 베루코마이크로비아 종, 월리넬라 종 (예를 들어, 월리넬라 숙시노게네스), 및/또는 게놈 데이터베이스의 생물정보학 조사에 기술된 종들, 가령, Makarova, Kira S., 등 "An updated evolutionary classification of CRISPR-Cas systems." Nature Reviews Microbiology 13.11 (2015): 722 and Koonin, Eugene V., Kira S. Makarova, and Feng Zhang. "Diversity, classification and evolution of CRISPR-Cas systems." Current opinion in microbiology 37 (2017): 67-78에 개시된 것들이 포함되며, 이들 각각은 본원에 그 전문이 참고로 포함된다.
일부 구체예에서, 조작된 Cas9 단백질은 스트렙토코쿠스 피오게네스로부터 유래할 수 있다. 일부 구체예에서, 조작된 Cas9 단백질은 스트렙토코쿠스 써모필루스로부터 유래할 수 있다. 일부 구체예에서, 조작된 Cas9 단백질은 네이세리아 메닝기티디스로부터 유래할 수 있다. 일부 구체예에서, 조작된 Cas9 단백질은 스타필로코쿠스 아우레우스로부터 유래할 수 있다. 일부 구체예에서, 조작된 Cas9 단백질은 캄필로박터 제주니로부터 유래할 수 있다.
야생형 Cas9 단백질은 두 개의 뉴클레아제 도메인, 즉, RuvC 및 HNH 도메인을 포함하며, 이들 각각은 이중 가닥 서열의 한 가닥을 절단한다. Cas9 단백질은 또한 가이드 RNA (예컨대, REC1, REC2) 또는 RNA/DNA 이종이중체 (예컨대, REC3)와 상호작용하는 REC 도메인들, 및 프로토스페이서-인접 모티프 (PAM)와 상호작용하는 도메인 (즉, PAM-상호작용 도메인)을 포함한다.
Cas9 단백질은 Cas9 단백질은 변경된 활성, 특이성 및/또는 안정성을 가지도록 하는 하나 이상의 변형 (즉, 하나 이상의 아미노산의 치환, 하나 이상의 아미노산의 결실, 하나 이상의 아미노산의 삽입)을 포함하도록 조작될 수 있다.
예를 들어, Cas9 단백질은 뉴클레아제 도메인 중 하나 또는 둘 모두를 비활성화하기 위한 하나 이상의 돌연변이 및/또는 결실에 의해 조작 될 수 있다. 하나의 뉴클레아제 도메인의 비활성화는 이중 가닥 서열의 한 가닥을 절단하는 Cas9 단백질 (즉, Cas9 니카아제)을 생성한다. RuvC 도메인은 D10A, D8A, E762A 및/또는 D986A와 같은 돌연변이에 의해 비활성화 될 수 있으며, HNH 도메인은 H840A, H559A, N854A, N856A 및/또는 N863A와 같은 돌연변이에 의해 비활성화 될 수 있다 (스트렙토코쿠스 피오게네스 Cas9, SpyCas9의 넘버링 시스템 참고). 두 뉴클레아제 도메인의 비활성화는 절단 활성이 없는 Cas9 단백질을 생성한다 (즉, 촉매적으로 비활성이거나 죽은 Cas9).
Cas9 단백질은 또한, 개선된 표적화 특이성, 개선된 충실도, 변경된 PAM 특이성, 감소된 오프-타겟 효과 및/또는 증가된 안정성을 가지도록 하는 하나 이상의 아미노산 치환, 결실 및/또는 삽입에 의해 조작될 수 있다. 표적화 특이성을 개선하고, 충실도를 개선하고, 및/또는 오프-타겟 효과를 감소시키는 하나 이상의 돌연변이의 비-제한적인 예는 N497A, R661A, Q695A, K810A, K848A, K855A, Q926A, K1003A, R1060A 및/또는 D1135E를 포함한다 (SpyCas9의 넘버링 체계 참고).
대안적인 구체예들에서, Cas 단백질은 유형 I CRISPR/Cas 시스템으로부터 유래할 수 있다. 일부 구체예들에서, Cas 단백질은 유형-I CRISPR/Cas 시스템의 캐스케이드 복합체의 구성성분일 수 있다. 예를 들어, Cas 단백질은 Cas3 단백질일 수 있다. 일부 구체예들에서, Cas 단백질은 유형 III CRISPR/Cas 시스템으로부터 유래할 수 있다. 일부 구체예들에서, Cas 단백질은 유형 IV CRISPR/Cas 시스템으로부터 유래할 수 있다. 일부 구체예들에서, Cas 단백질은 유형 V CRISPR/Cas 시스템으로부터 유래할 수 있다. 일부 구체예들에서, Cas 단백질은 유형 VI CRISPR/Cas 시스템으로부터 유래할 수 있다. 일부 구체예들에서, Cas 단백질은 RNA 절단 활성을 가질 수 있다. 다양한 구체예들에서, Cas 단백질은 Cas9, Cas12a (일명 Cpf1), Cas12b, Cas12c, Cas12d, Cas12e (일명 CasX), Cas13a, 또는 Cas13b로 분류될 수 있다.
(i) 이종 도메인
Cas9 단백질은 하나 이상의 이종 도메인을 포함하도록 조작 될 수 있다, 즉, Cas9는 하나 이상의 이종 도메인에 융합된다. 둘 이상의 이종 도메인이 Cas9와 융합되는 상황에서, 둘 이상의 이종 도메인은 동일하거나 다를 수 있다. 하나 이상의 이종 도메인은 N 말단 단부, C 말단 단부, 내부 위치 또는 이들의 조합에 융합 될 수 있다. 융합은 화학적 결합을 통한 직접 일 수 있으며, 또는 연결은 하나 이상의 링커를 통한 간접 일 수 있다.
특정 바람직한 구체예에서, 본원에 기재된 조작된 Cas9 단백질은 하나 이상의 핵 위치화 신호(NLS)를 포함한다. 핵 위치화 신호의 비-제한적 예들은 PKKKRKV (서열 번호: 1), PKKKRRV (서열 번호: 2), KRPAATKKAGQAKKKK (서열 번호: 3), YGRKKRRQRRR (서열 번호: 4), RKKRRQRRR (서열 번호: 5), PAAKRVKLD (서열 번호: 6), RQRRNELKRSP (서열 번호: 7), VSRKRPRP (서열 번호: 8), PPKKARED (서열 번호: 9), PQPKKKPL (서열 번호: 10), SALIKKKKKMAP (서열 번호: 11), PKQKKRK (서열 번호: 12), RKLKKKIKKL (서열 번호: 13), REKKKFLKRR (서열 번호: 14), KRKGDEVDGVDEVAKKKSKK (서열 번호: 15), RKCLQAGMNLEARKTKK (서열 번호: 16), NQSSNFGPMKGGNFGGRSSGPYGGGGQYFAKPRNQGGY (서열 번호: 17), 및 RMRIZFKNKGKDTAELRRRRVEVSVELRKAKKDEQILKRRNV (서열 번호: 18)를 포함한다.
한 특정 구체예에서, 핵 위치화 신호는 PKKKRKV (서열 번호: 1) 및 PAAKRVKLD (서열 번호: 6)에서 선택된다. 또 다른 특정 구체예에서, 조작된 Cas9 단백질은 PKKKRKV (서열 번호: 1) 및 PAAKRVKLD (서열 번호: 6)를 모두 포함한다. 또 다른 특정 구체예에서, 조작된 Cas9 단백질은 적어도 2개의 PKKKRKV (서열 번호: 1) 및 적어도 하나의 PAAKRVKLD (서열 번호: 6)를 포함한다. 또 다른 특정 구체예에서, 조작된 Cas9 단백질은 2개의 PKKKRKV (서열 번호: 1) 및 하나의 PAAKRVKLD (서열 번호: 6)를 포함한다.
이들 및 다른 바람직한 실시형태에서, 조작된 Cas9 단백질은 하나 이상의 마커 도메인을 포함한다. 마커 도메인은 형광 단백질 및 정제 또는 에피토프 태그를 포함한다. 적합한 형광 단백질들에는, 제한 없이, 녹색 형광 단백질 (예컨대, GFP, eGFP, GFP-2, tagGFP, turboGFP, Emerald, Azami Green, Monomeric Azami Green, CopGFP, AceGFP, ZsGreen1), 황색 형광 단백질 (예컨대, YFP, EYFP, Citrine, Venus, YPet, PhiYFP, ZsYellow1), 청색 형광 단백질 (예컨대, BFP, EBFP, EBFP2, Azurite, mKalama1, GFPuv, Sapphire, T-sapphire), 시안 형광 단백질 (예컨대, ECFP, Cerulean, CyPet, AmCyan1, Midoriishi-Cyan), 적색 형광 단백질 (예컨대, mKate, mKate2, mPlum, DsRed 단량체, mCherry, mRFP1, DsRed-Express, DsRed2, DsRed-Monomer, HcRed-Tandem, HcRed1, AsRed2, eqFP611, mRasberry, mStrawberry, Jred), 오렌지색 형광 단백질 (예컨대, mOrange, mKO, Kusabira-Orange, Monomeric Kusabira-Orange, mTangerine, tdTomato), 및 이의 조합이 포함된다. 마커 도메인은 하나 이상의 형광 단백질 (예컨대, Suntag)의 일렬 반복을 포함 할 수 있다.
한 구체예에서, 마커 단백질은 다음에서 선택된다:
적합한 정제 또는 에피토프 태그의 비-제한적 예들에는 6xHis (서열 번호: 22), FLAG® (예를 들어, 서열 번호: 21), HA, GST, Myc, SAM, 등이 포함된다. CRISPR 복합체의 검출 또는 농축을 용이하게 하는 이종 융합의 비-제한적 예들은 스트렙타비딘 (Kipriyanov 외, 인간 Antibodies, 1995, 6(3):93-101.), 아비딘 (Airenne 외, Biomolecular Engineering, 1999, 16(1-4):87-92), 아비딘의 단량체 형태 (Laitinen 외, Journal of Biological Chemistry, 2003, 278(6):4010-4014), 재조합 생산 동안 비오티닐화를 용이하게 하는 펩티드 태그 (Cull 외, Methods in Enzymology, 2000, 326:430-440)를 포함한다.
핵 위치화 신호(들) 및 마커 단백질(들) 이외에도, 다양한 구체예들에서 조작된 Cas9 단백질은 또한 하나 이상의 이종 도메인, 예를 들어, 세포-침투 도메인, 마커 도메인, 염색질 파괴 도메인, 후성 변형 도메인 (예를 들어, 시티딘 탈아미노효소 도메인, 히스톤 아세틸트랜스퍼라제 도메인 등), 전사 조절 도메인, RNA 압타머 결합 도메인, 또는 비-Cas9 뉴클레아제 도메인을 포함할 수 있다.
일부 구체예에서, 하나 이상의 이종 도메인은 세포-침투 도메인 일 수 있다. 적합한 세포-침투 도메인들의 예들은, 제한없이, GRKKRRQRRRPPQPKKKRKV (서열 번호: 23), PLSSIFSRIGDPPKKKRKV (서열 번호: 24), GALFLGWLGAAGSTMGAPKKKRKV (서열 번호: 25), GALFLGFLGAAGSTMGAWSQPKKKRKV (서열 번호: 26), KETWWETWWTEWSQPKKKRKV (서열 번호: 27), YARAAARQARA (서열 번호: 28), THRLPRRRRRR (서열 번호: 29), GGRRARRRRRR (서열 번호: 30), RRQRRTSKLMKR (서열 번호: 31), GWTLNSAGYLLGKINLKALAALAKKIL (서열 번호: 32), KALAWEAKLAKALAKALAKHLAKALAKALKCEA (서열 번호: 33), 및 RQIKIWFQNRRMKWKK (서열 번호: 34)를 포함한다.
또한 다른 구체예들에서, 하나 이상의 이종 도메인은 염색질 조절 모티프 (CMM) 일 수 있다. CMM들의 비-제한적 예들은 고 이동성 그룹 (HMG) 단백질들 (예컨대, HMGB1, HMGB2, HMGB3, HMGN1, HMGN2, HMGN3a, HMGN3b, HMGN4, 및 HMGN5 단백질), 히스톤 H1 변이체들 (예컨대, 히스톤 H1.0, H1.1, H1.2, H1.3, H1.4, H1.5, H1.6, H1.7, H1.8, H1.9, 및 H.1.10)의 중심 구상 도메인, 또는 염색질 재형성 복합체의 DNA 결합 도메인 (예컨대, SWI/SNF (스위치/수크로스 비-발효성, SWItch/Sucrose Non-Fermentable), ISWI (모방 스위치, Imitation SWItch), CHD (크로모도메인-헬리카제-DNA 결합, Chromo도메인-헬리카제-DNA binding), Mi-2/NuRD (뉴클레오솜 재형성 및 탈아세틸화효소, Nucleosome Remodeling and 탈아세틸화효소), INO80, SWR1, 및 RSC 복합체에서 유래한 뉴클레오솜 상호작용 펩티드를 포함한다. 다른 구체예들에서, CMM들은 또한 토포아이소머라제, 헬리카제, 또는 바이러스 단백질에서 유래될 수 있다. CMM의 공급원은 변화할 수 있으며 변화할 것이다. CMM들은 인간, 동물 (즉, 척추동물 및 무척추동물), 식물, 조류, 또는 효모에서 유래할 수 있다. 특정 CMM의 비 제한적인 예가 아래 표에 나열되어 있다. 당업자는 다른 종들의 상동체 및/또는 그 내부의 관련 융합 모티프를 용이하게 확인할 수 있다.
또 다른 구체예에서, 하나 이상의 이종 도메인은 후성 변형 도메인 일 수 있다. 적합한 후성 변형 도메인의 비-제한적 예들은 DNA 탈아미노화 (예컨대, 시티딘 탈아미노효소, 아데노신 탈아미노효소, 구아닌 탈아미노효소), DNA 메틸전이효소 활성 (예컨대, 시토신 메틸전이효소), DNA 탈메틸효소 활성, DNA 아미노화, DNA 산화 활성, DNA 헬리카제 활성, 히스티딘 아세틸전이효소 (HAT) 활성 (예컨대, E1A 결합 단백질 p300에서 유래한 HAT 도메인), 히스티딘 탈아세틸화효소 활성, 히스티딘 메틸전이효소 활성, 히스티딘 탈메틸효소 활성, 히스티딘 키나제 활성, 히스티딘 포스파타제 활성, 히스티딘 유비퀴틴 리가제 활성, 히스티딘 틸유비퀴틴화 활성, 히스티딘 아데닐화 활성, 히스티딘 탈아데닐화 활성, 히스티딘 SUMO일화 활성, 히스티딘 탈SUMOy일화 활성, 히스티딘 리보실화 활성, 히스티딘 탈리보실화 (deribosylation) 활성, 히스티딘 미리스토일화 활성, 히스티딘 탈미리스토일화 활성, 히스티딘 시트룰린화 활성, 히스티딘 알킬화 활성, 히스티딘 탈알킬화 활성, 또는 히스티딘 산화 활성을 가지진 것을 포함한다. 특정 구체예들에서, 후성 변형 도메인은 시티딘 탈아미노효소 활성, 아데노신 탈아미노효소 활성, 히스톤 아세틸전이효소 활성, 또는 DNA 메틸전이효소 활성을 포함할 수 있다.
다른 구체예에서, 하나 이상의 이종 도메인은 전사 조절 도메인 (즉, 전사 활성화 도메인 또는 전사 억제 도메인) 일 수 있다. 적합한 전사 활성화 도메인은 제한없이 단순 헤르페스 바이러스 VP16 도메인, VP64 (즉, VP16의 4개의 탠덤 사본), VP160 (즉, VP16의 10개의 탠덤 사본), NFκp65 활성화 도메인 (p65), 엡스타인-바 바이러스 R 트랜스활성화인자 (Rta) 도메인, VPR (즉, VP64+p65+Rta), p300-의존성 전사 활성화 도메인, p53 활성화 도메인 1 및 2, 열 충격 인자 1 (HSF1) 활성화 도메인, Smad4 활성화 도메인 (SAD), cAMP 반응 요소 결합 단백질 (CREB) 활성화 도메인, E2A 활성화 도메인, 활성화된 T 세포의 핵 인자 (NFAT) 활성화 도메인, 또는 이들의 조합을 포함한다. 적합한 전사 억제자 도메인의 비 제한적인 예는 크루펠-관련 박스 (KRAB) 억제자 도메인, Mxi 억제자 도메인, 유도성 cAMP 조기 억제자 (ICER) 도메인, YY1 글리신 풍부 억제자 도메인, Sp1-유사 억제자, E(spl) 억제자를 포함하며, Iκ억제자, Sin3 억제자, 메틸-CpG 결합 단백질 2 (MeCP2) 억제자, 또는 이들의 조합을 포함한다. 전사 활성화 또는 전사 억제인자 도메인은 Cas9 단백질에 유전적으로 융합되거나 비공유 단백질-단백질, 단백질-RNA, 또는 단백질-DNA 상호작용에 의해 결합될 수 있다.
추가 구체예에서, 하나 이상의 이종 도메인은 RNA 압타머 결합 도메인 일 수 있다 (Konermann 외, Nature, 2015, 517(7536):583-588; Zalatan 외, Cell, 2015, 160(1-2):339-50). 적합한 RNA 압타머 단백질 도메인의 예들에는 MS2 외피 단백질 (MCP), PP7 박테리오파지 외피 단백질 (PCP), Mu 박테리오파지 Com 단백질, 람다 박테리오파지 N22 단백질, 스템-루프 결합 단백질 (SLBP), 취약 X 정신 지체 증후군-관련 단백질 1 (FXR1), 박테리오파지로부터 유래한 단백질, 가령, AP205, BZ13, f1, f2, fd, fr, ID2, JP34/GA, JP501, JP34, JP500, KU1, M11, M12, MX1, NL95, PP7, φCb5, φCb8r, φCb12r, φCb23r, Qβ, R17, SP-β, TW18, TW19 및 VK, 이의 단편, 또는 이의 유도체가 포함된다.
또 다른 구체예에서, 하나 이상의 이종 도메인은 비-Cas9 뉴클레아제 도메인 일 수 있다. 적합한 뉴클레아제 도메인은 임의의 엔도뉴클레아제 또는 엑소뉴클레아제로부터 얻을 수 있다. 뉴클레아제 도메인이 유도될 수 있는 엔도뉴클레아제의 비-제한적 예들은, 제한 엔도뉴클레아제 및 호밍 엔도뉴클레아제를 포함하나 이에 제한되는 것은 아니다. 일부 구체예들에서, 뉴클레아제 도메인은 유형 II-S 제한 엔도뉴클레아제로부터 유래될 수 있다. 유형 II-S 엔도뉴클레아제는 인식/결합 부위로부터 전형적으로 여러 개 염기쌍 떨어져있는 부위들의 DNA를 절단하고, 그리하여 분리가능한 결합 및 절단 도메인을 가진다. 이들 효소들은 일반적으로 엇갈린 위치들에서 DNA 각 가닥을 절단하기 위해 일시적으로 결합하여 이량체를 형성하는 단량체이다. 적합한 유형 II-S 엔도뉴클레아제의 비-제한적 예들에는 BfiI, BpmI, BsaI, BsgI, BsmBI, BsmI, BspMI, FokI, MboII, 및 SapI가 포함된다. 일부 구체예들에서, 뉴클레아제 도메인은 FokI 뉴클레아제 도메인 또는 이의 유도체일 수 있다. 유형 II-S 뉴클레아제 도메인은 2개의 상이한 뉴클레아제 도메인의 이량체화를 용이하게 하도록 변형될 수 있다. 예를 들면, FokI의 절단 도메인은 특정 아미노산 잔기들을 돌연변이시킴으로써 변형될 수 있다. 비-제한적 예로서, FokI 뉴클레아제 도메인의 위치 446, 447, 479, 483, 484, 486, 487, 490, 491, 496, 498, 499, 500, 531, 534, 537, 및 538의 아미노산 잔기들은 변형을 위한 표적이다. 특정 구체예들에서, FokI 뉴클레아제 도메인은 Q486E, I499L, 및/또는 N496D 돌연변이들을 포함하는 제 1 FokI 절반-도메인, 및 E490K, I538K, 및/또는 H537R 돌연변이들을 포함하는 제 2 FokI 절반-도메인을 포함할 수 있다.
하나 이상의 이종 도메인은 하나 이상의 화학적 결합 (예를 들어, 공유 결합)을 통해 Cas9 단백질에 직접 연결될 수 있거나, 하나 이상의 이종 도메인이 하나 이상의 링커를 통해 Cas9 단백질에 간접적으로 연결될 수 있다.
링커는 최소한 하나의 공유 결합에 의해 하나 이상의 다른 화학적 작용기를 연결시키는 화학적 작용기이다. 적합한 링커들에는 아미노산, 펩티드, 뉴클레오티드, 핵산, 유기 링커 분자들 (예컨대, 말레이미드 유도체, N-에톡시벤질이미다졸, 바이페닐-3,4′트라이카르복시산, p-아미노벤질옥시카르보닐, 등), 다이설파이드 링커, 및 폴리머 링커 (예컨대, PEG)가 포함된다. 링커는 알킬렌, 알켄일렌, 알킨일렌, 알킬, 알켄일, 알킨일, 알콕시, 아릴, 헤테로아릴, 아랄킬, 아랄켄일, 아랄킨일 등을 비롯한 (그러나 이에 제한되는 것은 아님) 하나 이상의 스페이싱 작용기들을 포함할 수 있다. 링커는 중성일 수 있거나, 양 또는 음 전하를 지닐 수 있다. 추가적으로, 링커는 절단가능할 수 있어서, 링커를 또 다른 화학적 작용기에 연결시키는 링커의 공유 결합이, pH, 온도, 염 농도, 빛, 촉매 또는 효소를 포함한 특정 조건하에서 파열되거나 절단될 수 있다. 일부 구체예들에서, 링커는 펩티드 링커일 수 있다. 펩티드 링커는 가요성 아미노산 링커 (예컨대, 소형, 비-극성 또는 극성 아미노산을 포함) 일 수 있다.
특정 구체예에서, 링커는 다음에서 선택된다:
가요성 링커의 다른 비-제한적 예들은 LEGGGS (서열 번호: 37), TGSG (서열 번호: 38), GGSGGGSG (서열 번호: 39), (GGGGS)1-4 (서열 번호: 40), 및 (Gly)6-8 (서열 번호: 41)을 포함한다. 대안적으로, 펩티드 링커는 단단한 아미노산 링커일 수 있다. 이러한 링커들은 (EAAAK)1-4 (서열 번호: 42), A(EAAAK)2-5A (서열 번호: 43), PAPAP (서열 번호: 44), 및 (AP)6-8 (서열 번호: 45)을 포함한다. 적합한 링커들의 또 다른 예는 해당 분야에 잘 공지되어 있으며 링커들을 설계하는 프로그램들은 용이하게 이용가능하다 (Crasto 외., Protein Eng., 2000, 13(5):309-312).
일부 구체예에서, 조작된 Cas9 단백질은 무세포 시스템, 박테리아 세포 또는 진핵 세포에서 재조합적으로 생산 될 수 있고 표준 정제 수단을 사용하여 정제될 수 있다. 다른 구체예들에서, 조작된 Cas9 단백질은 조작된 Cas9 단백질을 인코딩하는 핵산으로부터 관심 진핵 세포의 생체내에서 생산된다 (아래 섹션 (II) 참고).
조작된 Cas9 단백질이 뉴클레아제 또는 니카아제 활성을 포함하는 구체예에서, 조작된 Cas9 단백질은 적어도 하나의 세포-침투 도메인, 뿐만 아니라 적어도 하나의 염색질 파괴 도메인을 추가로 포함 할 수 있다. 조작된 Cas9 단백질이 후성 변형 도메인에 연결되는 구체예에서, 조작된 Cas9 단백질은 적어도 하나의 세포-침투 도메인, 뿐만 아니라 적어도 하나의 염색질 파괴 도메인을 추가로 포함 할 수 있다. 또한, 조작된 Cas9 단백질이 전사 조절 도메인에 연결되는 구체예에서, 조작된 Cas9 단백질은 적어도 하나의 세포-침투 도메인, 뿐만 아니라 적어도 하나의 염색질 파괴 도메인 및/또는 적어도 하나의 RNA 압타머 결합 도메인을 추가로 포함 할 수 있다.
다양한 융합 단백질 성분은 N-말단에서 C-말단까지 임의의 순서로 조합될 수 있다. 예를 들어, A가 마커 단백질을 나타내고, B가 핵 위치화 신호를 나타내고, C가 Cas9 단백질을 나타내는 경우, 융합 단백질은 다음 방식으로 N-말단에서 C-말단으로 배열될 수 있다: A-B-C; A-C-B; B-A-C; B-C-A; C-A-B; 또는 C-B-A, 이때 링커 (“-L-”)는 임의의 2개 항목들 사이에 배치될 수 있다 (예를 들어, A-L-B-C; A-B-L-C; A-L-B-L-C; 등).
(b) 조작된 가이드 RNA
조작된 가이드 RNA는 특정 조작된 Cas9 단백질과 복합체를 형성하도록 설계된다. 가이드 RNA는 (i) 표적 서열과 혼성화하는, 5' 단부에 가이드 서열을 내포하는 CRISPR RNA (crRNA) 및 (ii) Cas9 단백질을 동원하는 트랜스작용 crRNA (tracrRNA) 서열을 포함한다. 각 가이드 RNA의 crRNA 가이드 서열은 상이하다 (즉, 서열 특이적이다). tracrRNA 서열은 일반적으로 특정 박테리아 종들의 Cas9 단백질과 복합체를 형성하도록 설계된 가이드 RNA에서 동일하다.
crRNA 가이드 서열은 이중 가닥 서열에서 표적 서열 (즉, 프로토스페이서)과 혼성화하도록 설계된다. 일반적으로, crRNA와 표적 서열 간의 상보성은 최소한 80%, 최소한 85%, 최소한 90%, 최소한 95%, 또는 최소한 99%이다. 특정 구체예들에서, 상보성은 완전하다 (즉, 100%). 다양한 구체예들에서, crRNA 가이드 서열의 길이는 약 15개 뉴클레오티드 내지 약 25개 뉴클레오티드 범위일 수 있다. 예를 들면, crRNA 가이드 서열은 약 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 또는 25개 뉴클레오티드 길이일 수 있다. 특정 구체예들에서, crRNA는 약 19, 20, 또는 21개 뉴클레오티드 길이이다. 한 구체예에서, crRNA 가이드 서열은 20개 뉴클레오티드의 길이를 가진다.
가이드 RNA는 Cas9 단백질과 상호작용하는 하나 이상의 스템 루프 구조를 형성하는 반복 서열 및 단일 가닥으로 남아있는 3' 서열을 포함한다. 각 루프와 스템의 길이는 다를 수 있습니다. 예를 들면, 루프는 약 3 내지 약 10개 뉴클레오티드 길이 범위일 수 있으며, 스템은 약 6개 내지 약 20개 염기쌍 길이 범위일 수 있다. 스템은 하나 이상의 1 내지 약 10개 뉴클레오티드 팽창부를 포함할 수 있다. 단일 가닥 3' 영역의 길이는 다를 수 있다. 조작된 가이드 RNA에서 tracrRNA 서열은 일반적으로 관심 박테리아 종들에서의 야생형 tracrRNA의 코딩 서열에 기초한다. 야생형 서열은 2차 구조 형성을 촉진, 2차 구조 안정성을 증가, 진핵 세포에서 발현을 촉진하도록 변형 될 수 있다. 예를 들면, 하나 이상의 뉴클레오티드 변화가 가이드 RNA 코딩 서열에 도입될 수 있다 (아래 실시예 3 참고). tracrRNA 서열은 약 50개 뉴클레오티드 내지 약 300개 뉴클레오티드 길이 범위일 수 있다. 다양한 구체예들에서, tracrRNA는 약 50 내지 약 90개 뉴클레오티드, 약 90 내지 약 110개 뉴클레오티드, 약 110 내지 약 130개 뉴클레오티드, 약 130 내지 약 150개 뉴클레오티드, 약 150 내지 약 170개 뉴클레오티드, 약 170 내지 약 200개 뉴클레오티드, 약 200 내지 약 250개 뉴클레오티드, 또는 약 250 내지 약 300개 뉴클레오티드 길이 범위일 수 있다.
일반적으로, 조작된 가이드 RNA는 단일 분자 (즉, 단일 가이드 RNA 또는 sgRNA)이며, 이 때 crRNA 서열은 tracrRNA 서열에 연결된다. 그러나 일부 구체예들에서, 조작된 가이드 RNA는 2개의 별도 분자들 일 수 있다. 제 1 분자는 제 2 분자의 5' 단부와 염기쌍을 이룰 수 있는 3' 서열 (약 6 내지 약 20개 뉴클레오티드를 포함)을 함유하는 crRNA를 포함하고, 이 때 제 2 분자는 제 1 분자의 3' 단부와 염기쌍을 이룰 수 있는 5' 서열 (약 6 내지 약 20개 뉴클레오티드를 포함)을 함유하는 tracrRNA를 포함한다.
일부 구체예들에서, 조작된 가이드 RNA의 tracrRNA 서열은 하나 이상의 압타머 서열들을 포함하도록 변형될 수 있다 (Konermann 외., Nature, 2015, 517(7536):583-588; Zalatan 외., Cell, 2015, 160(1-2):339-50). 적합한 압타머 서열들은 MCP, PCP, Com, SLBP, FXR1, AP205, BZ13, f1, f2, fd, fr, ID2, JP34/GA, JP501, JP34, JP500, KU1, M11, M12, MX1, NL95, PP7, φCb5, φCb8r, φCb12r, φCb23r, Qβ, R17, SP-β, TW18, TW19, VK, 이의 단편, 또는 이의 유도체에서 선택된 압타머 단백질들에 결합하는 서열들을 포함한다. 당업자는 압타머 길이가 달라질 수 있음을 이해한다.
다른 구체예들에서, 가이드 RNA는 최소한 하나의 탐지가능한 표지를 추가로 포함할 수 있다. 탐지가능한 표지는 형광단 (예컨대, FAM, TMR, Cy3, Cy5, 텍사스 레드, 오레곤 그린, 알렉사 플루오르, 할로 태그, 또는 적합한 형광 염료), 탐지 태그 (예컨대, 비오틴, 디곡시제닌, 등), 양자 점, 또는 금 입자들 일 수 있다.
가이드 RNA는 표준 리보뉴클레오티드 및/또는 변형된 리보뉴클레오티드를 포함 할 수 있다. 일부 구체예에서, 가이드 RNA는 표준 또는 변형된 데옥시리보뉴클레오티드를 포함할 수 있다. 가이드 RNA가 효소적으로 합성되는 (즉, 생체내 또는 시험관내) 구체예에서, 가이드 RNA는 일반적으로 표준 리보뉴클레오티드를 포함한다. 가이드 RNA가 화학적으로 합성되는 구체예들에서, 가이드 RNA는 표준 또는 변형된 리보뉴클레오티드 및/또는 데옥시리보뉴클레오티드를 포함할 수 있다. 변형된 리보뉴클레오티드 및/또는 데옥시리보뉴클레오티드는 염기 변형 (예컨대, 슈도우리딘, 2-티오우리딘, N6-메틸아데노신 등) 및/또는 당 변형 (예컨대, 2'-O-메틸, 2'-플루오로, 2'-아미노, 잠금 핵산 (LNA) 등)을 포함한다. 가이드 RNA의 골격은 또한 포스포로티오에이트 링키지, 보라노포스페이트 링키지 또는 펩티드 핵산을 포함하도록 변형 될 수 있다.
(c) PAM 서열
일부 구체예에서, 표적 서열은 CRISPR/Cas9 복합체에 의해 인식되는 짧은 서열인 프로토스페이서 인접 모티프(PAM)에 인접할 수 있다. 일부 구체예에서, PAM은 표적 서열의 3' 말단의 1, 2, 3 또는 4개의 뉴클레오티드에 인접하거나 그 내부에 존재할 수 있다. PAM의 길이와 서열은 사용된 Cas9 단백질에 따라 달라질 수 있다. 예를 들어, PAM은 Ran 등, Nature, 520: 186-191(2015) 본원에 참고로 포함됨)의 도 1에 도시된 것들을 포함하는 특정 Cas9 단백질 또는 Cas9 오르토로그에 대한 공통 또는 특정 PAM 서열로부터 선택될 수 있다. 일부 구체예들에서, PAM은 2, 3, 4, 5, 6, 7, 8, 9, 또는 10개 뉴클레오티드 길이를 포함할 수 있다. 비-제한적인 예시적 PAM 서열들은 NGG, NGGNG, NG, NAAAAN, NNAAAAW, NNNNACA, GNNNCNNA, 및 NNNNGATT를 포함한다 (이때 N은 임의의 뉴클레오티드로 정의되고, W는 A 또는 T로 정의됨). 일부 구체예들에서, PAM 서열은 NGG 일 수 있다. 일부 구체예들에서, PAM 서열은 NGGNG 일 수 있다. 일부 구체예들에서, PAM 서열은 NNAAAAW 일 수 있다.
상이한 CRISPR 단백질이 상이한 PAM 서열을 인식한다는 것이 이해될 것이다. 예를 들어, Cas9 단백질들에 대한 PAM 서열들은 5'-NGG, 5'-NGGNG, 5'-NNAGAAW, 5'-NNNNGATT, 5-NNNNRYAC, 5'-NNNNCAAA, 5'-NGAAA, 5'-NNAAT, 5'-NNNRTA, 5'-NNGG, 5'-NNNRTA, 5'-MMACCA, 5'-NNNNGRY, 5'-NRGNK, 5'-GGGRG, 5'-NNAMMMC, 및 5'-NNG를 포함하고, 및 Cas12a 단백질들에 대한 PAM 서열들은 5'-TTN 및 5'-TTTV를 포함하고, 이때 N은 임의의 뉴클레오티드로 정의되고, R은 G 또는 A로 정의되고, W는 A 또는 T로 정의되고, Y는 C 또는 T로 정의되고, 그리고 V는 A, C, 또는 G로 정의된다. 일반적으로, Cas9 PAM들은 표적 서열의 3'에 위치하고, 그리고 Cas12a PAM들은 표적 서열의 5'에 위치한다. 다양한 PAM 서열 및 이를 인식하는 CRISPR 단백질은 당업계,예를 들어, 미국 특허출원 공개공보 2019/0249200; Leenay, Ryan T., 등 "Identifying and visualizing functional PAM diversity across CRISPR-Cas systems." Molecular cell 62.1 (2016): 137-147; and Kleinstiver, Benjamin P., 등 "Engineered CRISPR-Cas9 nucleases with altered PAM specificities." Nature 523.7561 (2015): 481에 공지되어 있으며, 이들 각각은 본원에 그 전문이 참고로 포함된다.
추가적으로 또는 대안적으로, 본 명세서에 개시된 조작된 Cas9 시스템 각각에 대한 PAM이 하기에 제시되어 있다.
예를 들어, 미국 특허출원 공개공보 제 2019/0249200 (본원에 그 전문이 참고로 포함됨)을 참조하라.
(II)
핵산
본 발명의 또 다른 양상은 섹션 (I)에 상기 기재된 조작된 Cas9 시스템을 인코딩하는 핵산을 제공한다. 상기 시스템은 단일 핵산 또는 복수의 핵산들에 의해 인코딩 될 수 있다. 핵산은 DNA 또는 RNA, 선형 또는 원형, 단일-가닥 또는 이중-가닥 일 수 있다. RNA 또는 DNA는 관심 진핵 세포에서 단백질로의 효율적인 번역을 위해 코돈 최적화될 수 있다. 코돈 최적화 프로그램은 프리웨어로서 또는 상업적 공급업체들로부터 이용가능하다.
일부 구체예들에서, 핵산은 서열 번호: 48, 49, 또는 50의 아미노산 서열에 적어도 약 75%, 적어도 약 80%, 적어도 약 85%, 적어도 약 90%, 적어도 약 95% 또는 적어도 약 99% 서열 동일성을 가지는 단백질을 인코딩한다. 특정 구체예들에서, 조작된 Cas9 단백질을 인코딩하는 핵산은 서열 번호: 48, 49, 또는 50의 DNA 서열에 적어도 약 75%, 적어도 약 80%, 적어도 약 85%, 적어도 약 90%, 적어도 약 95% 또는 적어도 약 99% 서열 동일성을 가질 수 있다. 특정 구체예에서, 조작된 Cas9 단백질을 인코딩하는 DNA는 서열 번호: 48, 49, 또는 50의 DNA 서열을 가진다. 추가 구체예들에서, 핵산은 서열 번호: 48, 49, 또는 50의 아미노산 서열에 적어도 약 75%, 적어도 약 80%, 적어도 약 85%, 적어도 약 90%, 적어도 약 95% 또는 적어도 약 99% 서열 동일성을 가지는 단백질을 인코딩한다.
일부 구체예에서, 조작된 Cas9 단백질을 인코딩하는 핵산은 RNA일 수 있다. RNA는 시험관내에서 효소적으로 합성될 수 있다. 이를 위하여, 조작된 Cas9 단백질을 인코딩하는 DNA는 시험관내 RNA 합성을 위해 파지 RNA 중합효소에 의해 인식되는 프로모터 서열에 작동적으로 연결될 수 있다. 예를 들면, 프로모터 서열은 T7, T3, 또는 SP6 프로모터 서열 또는 T7, T3, 또는 SP6 프로모터 서열의 변이 일 수 있다. 조작된 단백질을 인코딩하는 DNA는 아래 상세히 설명된 벡터의 일부 일 수 있다. 이러한 구체예들에서, 시험관내-전사된 RNA는 정제, 캡핑, 및/또는 폴리아데닐화 될 수 있다. 다른 구체예들에서, 조작된 Cas9 단백질을 인코딩하는 RNA는 자가-복제 RNA의 일부 일 수 있다 (Yoshioka 외., Cell Stem Cell, 2013, 13:246-254). 자가-복제 RNA는 비감염성, 자가-복제 Venezuelan 말 뇌염 (VEE) 바이러스 RNA 레플리콘으로부터 유래될 수 있는데, 이는 제한된 세포 분열수를 위하여 자가-복제할 수 있는 양성-센스, 단일-가닥 RNA로서, 관심 단백질들을 코드화하기 위해 변형될 수 있다 (Yoshioka 외., Cell Stem Cell, 2013, 13:246-254).
다른 구체예에서, 조작된 Cas9 단백질을 인코딩하는 핵산은 DNA 일 수 있다. DNA 코딩 서열은 관심 세포에서 발현하기 위해 최소한 하나의 프로모터 제어 서열에 작동적으로 연결될 수 있다. 특정 구체예들에서, DNA 코딩 서열은 박테리아 (예컨대, 대장균) 세포 또는 진핵생물 (예컨대, 효모, 곤충, 또는 포유동물) 세포에서 조작된 Cas9 단백질의 발현을 위해 프로모터 서열에 작동적으로 연결될 수 있다. 적합한 박테리아 프로모터에는, 제한 없이, T7 프로모터, lac 오페론 프로모터, trp 프로모터, tac 프로모터 (이들은 trp과 lac 프로모터의 하이브리드임), 전술한 것들 중 어느 하나의 변이형, 및 전술한 것들의 임의의 조합이 포함된다. 적합한 진핵생물 프로모터의 비-제한적 예들에는 항시성의, 조절된 또는 세포- 또는 조직-특이적 프로모터가 포함된다. 적합한 진핵생물 항시적 프로모터 제어 서열에는, 거대세포바이러스 즉시 초기 프로모터 (CMV), 원숭이 바이러스 (SV40) 프로모터, 아데노바이러스 주요 후기 프로모터, 라우스 육종 바이러스 (RSV) 프로모터, 마우스 젖 종양 바이러스 (MMTV) 프로모터, 포스포글리세레이트 키나제 (PGK) 프로모터, 연장 인자 (ED1)-알파 프로모터, 유비퀴틴 프로모터, 액틴 프로모터, 튜불린 프로모터, 면역글로불린 프로모터, 이의 단편, 또는 전술한 것들의 임의의 조합이 포함되나, 이에 제한되는 것은 아니다. 적합한 진핵생물 조절된 프로모터 제어 서열의 예들에는 제한 없이 열 충격, 금속, 스테로이드, 항생제, 또는 알콜에 의해 조절되는 것들이 포함된다. 조직-특이적 프로모터의 비-제한적 예들에는 B29 프로모터, CD14 프로모터, CD43 프로모터, CD45 프로모터, CD68 프로모터, 데스민 프로모터, 엘라스타제-1 프로모터, 엔도글린 프로모터, 피브로넥틴 프로모터, Flt-1 프로모터, GFAP 프로모터, GPIIb 프로모터, ICAM-2 프로모터, INF-β 프로모터, Mb 프로모터, NphsI 프로모터, OG-2 프로모터, SP-B 프로모터, SYN1 프로모터, 및 WASP 프로모터가 포함된다. 프로모터 서열은 야생형 일 수 있거나 보다 효율적인 또는 효과적인 발현을 위해 변형될 수 있다. 일부 구체예들에서, DNA 코딩 서열은 또한 폴리아데닐화 신호 (예컨대, SV40 폴리A 신호, 소 성장 호르몬 (BGH) 폴리A 신호, 등) 및/또는 최소한 하나의 전사 종결 서열에 연결될 수 있다. 어떤 상황에서는 조작된 Cas9 단백질이 박테리아 또는 진핵 세포에서 정제 될 수 있다.
또 다른 구체예에서, 조작된 가이드 RNA는 DNA에 의해 인코딩 될 수 있다. 일부 예들에서, 조작된 가이드 RNA를 인코딩하는 DNA는 시험관내 RNA 합성을 위해 파지 RNA 중합효소에 의해 인식되는 프로모터 서열에 작동적으로 연결될 수 있다. 예를 들면, 프로모터 서열은 T7, T3, 또는 SP6 프로모터 서열 또는 T7, T3, 또는 SP6 프로모터 서열의 변이 일 수 있다. 다른 예에서, 조작된 가이드 RNA를 인코딩하는 DNA는 관심 진핵 세포에서의 발현을 위해 RNA 중합효소 III (Pol III)에 의해 인식되는 프로모터 서열에 작동가능하게 연결될 수 있다. 적합한 Pol III 프로모터의 예들에는, 포유동물 U6, U3, H1, 및 7SL RNA 프로모터가 포함되나, 이에 제한되는 것은 아니다.
다양한 구체예에서, 조작된 Cas9 단백질을 인코딩하는 핵산은 벡터에 존재할 수 있다. 일부 구체예에서, 벡터는 조작된 가이드 RNA를 인코딩하는 핵산을 추가로 포함 할 수 있다. 적합한 벡터들에는 플라스미드 벡터, 바이러스 벡터, 및 자가-복제 RNA가 포함된다 (Yoshioka 외., Cell Stem Cell, 2013, 13:246-254). 일부 구체예들에서, 복합체 또는 융합 단백질을 인코딩하는 핵산은 플라스미드 벡터에 존재할 수 있다. 적합한 플라스미드 벡터의 비-제한적 예들에는 pUC, pBR322, pET, pBluescript, 및 이의 변이체들이 포함된다. 다른 구체예들에서, 복합체 또는 융합 단백질을 인코딩하는 핵산은 바이러스 벡터의 일부 일 수 있다 (예컨대, 렌티바이러스 벡터, 아데노-연관 바이러스 벡터, 아데노바이러스 벡터, 등). 플라스미드 또는 바이러스 벡터는 또 다른 발현 제어 서열 (예컨대, 인핸서 서열, 코작 서열, 폴리아데닐화 서열, 전사 종결 서열, 등.), 선별 마커 서열 (예컨대, 항생제 내성 유전자), 복제 원점, 등을 포함할 수 있다. 벡터 및 이의 용도에 관한 추가 정보는 “Current Protocols in Molecular Biology” Ausubel 외., John Wiley & Sons, New York, 2003 or “Molecular Cloning: A Laboratory Manual” Sambrook & Russell, Cold Spring Harbor Press, Cold Spring Harbor, NY, 3rd edition, 2001에서 찾을 수 있다.
(III)
진핵 세포
본 발명의 또 다른 양상은 상기 섹션 (I)에 설명된 적어도 하나의 조작된 Cas9 시스템 및/또는 상기 섹션 (II)에 설명된 조작된 Cas9 단백질 및/또는 조작된 가이드 RNA를 인코딩하는 적어도 하나의 핵산을 포함하는 진핵 세포를 포함한다.
진핵 세포는 인간 세포, 비-인간 포유동물 세포, 비-포유동물 척추동물 세포, 무척추동물 세포, 식물 세포, 또는 단세포 진핵 유기체일 수 있다. 적합한 진핵 세포들의 예들은 상기 섹션 (IV)(c)에 설명된다. 진핵 세포는 시험관내, 세포외, 또는 세포내 일 수 있다.
예로서, 일부 구체예들에서, 진핵 세포 또는 진핵 세포 집단은 T 세포, CD8+ T 세포, CD8+ 나이브 T 세포, 중추 기억 T 세포, 효과기 기억 T 세포, CD4+ T 세포, 줄기 세포 기억 T 세포, 보조 T 세포, 조절 T 세포, 세포독성 T 세포, 자연 살해 T 세포, 조혈 줄기 세포, 장기 조혈 줄기 세포, 단기 조혈 줄기 세포, 다분화능 전구 세포, 계통 제한 전구 세포, 림프 전구 세포, 췌장 전구 세포, 내분비 전구 세포, 외분비 전구 세포, 골수 전구 세포, 일반 골수 전구 세포, 적혈구 전구 세포, 거핵 세포 적혈구 전구 세포, 단핵구 전구 세포, 내분비 전구 세포, 외분비 세포, 섬유아세포, 간모세포, 근모세포, 대식세포, 섬 베타 세포, 심근세포, 혈액 세포, 관 세포, 포상 세포, 알파 세포, 베타 세포, 델타 세포, PP 세포, 담관 세포, 망막 세포, 광수용기 세포, 간상 세포, 원추 세포, 망막 색소 상피 세포, 섬유주 세포, 와우 세포, 외유모 세포, 내유모 세포, 폐 상피 세포, 기관지 상피 세포, 폐포 상피 세포, 폐 상피 전구 세포, 횡문근 세포, 심장 근육 세포, 근육 위성 세포, 근세포, 뉴런, 뉴런 줄기 세포, 중간엽 줄기 세포, 유도 만능 줄기 세포(iPS) 세포 , 배아줄기세포, 단핵구, 거핵구, 호중구, 호산구, 호염기구, 비만세포, 망상적혈구, B 세포, 예를 들어, 전구 B 세포, 예를 들어, Pre B 세포, Pro B 세포, 기억 B 세포, 혈장 B 세포, 위장 상피 세포, 담도 상피 세포, 췌관 상피 세포, 장 줄기 세포, 간세포, 간 성상 세포, 쿠퍼 세포, 조골 세포, 파골 세포, 지방세포(예를 들어, 갈색 지방세포 또는 백색 지방세포), 전지방세포, 췌장 반응성 전구체 세포, 췌장 섬 세포, 췌장 베타 세포, 췌장 알파 세포, 췌장 델타 세포, 췌장 외분비 세포, 슈반 세포 또는 희돌기아교세포, 또는 이러한 세포의 집단이다.
(IV)
염색체 서열 변형 방법
본 발명의 추가 양상은 진핵 세포에서 염색체 서열을 변형하는 방법을 포함한다. 일반적으로, 이 방법은 섹션 (I)에서 상술한 적어도 하나의 조작된 Cas9 시스템 및/또는 섹션 (II)에서 상술한 상기 조작된 Cas9 시스템을 인코딩하는 적어도 하나의 핵산을 관심 진핵 세포에 도입하는 것을 포함한다.
조작된 Cas9 단백질이 뉴클레아제 또는 니카아제 활성을 포함하는 구체예에서, 염색체 서열 변형은 적어도 하나의 뉴클레오티드의 치환, 적어도 하나의 뉴클레오티드의 결실, 적어도 하나의 뉴클레오티드의 삽입을 포함 할 수 있다. 일부 구체예들에서, 상기 방법은 뉴클레아제 활성을 포함하는 하나의 조작된 Cas9 시스템 또는 니카아제 활성을 포함하고 공여체 폴리뉴클레오티드가 없는 2개의 조작된 Cas9 시스템을 진핵 세포에 도입하는 것을 포함하고, 이로써 조작된 Cas9 시스템 또는 시스템들은 염색체 서열의 표적 부위에 이중-가닥 절단을 도입하고 세포 DNA 복구 과정에 의한 이중-가닥 절단의 복구는 적어도 하나의 뉴클레오티드 변화 (즉, 삽입결실)를 도입함으로써, 염색체 서열을 비활성화 (즉, 유전자 녹아웃) 시킨다. 다른 구체예들에서, 상기 방법은 뉴클레아제 활성을 포함하는 하나의 조작된 Cas9 시스템 또는 니카아제 활성, 그리고 공여체 폴리뉴클레오티드를 포함하는 2개의 조작된 Cas9 시스템을 진핵 세포에 도입하는 것을 포함하고, 이로써 조작된 Cas9 시스템 또는 시스템들은 염색체 서열의 표적 부위에 이중-가닥 절단을 도입하고 세포 DNA 복구 과정에 의한 이중-가닥 절단의 복구는 공여체 폴리뉴클레오티드 서열을 염색체 서열의 표적 부위 내부에 삽입 또는 교환하는 결과를 가져온다.
조작된 Cas9 단백질이 후성 변형 활성 또는 전사 조절 활성을 포함하는 구체예들에서, 염색체 서열 변형은 염색체 서열의 표적 부위에서 또는 근방에서 적어도 하나의 뉴클레오티드의 전환, 표적 부위에서 또는 근방에서 적어도 하나의 뉴클레오티드의 변형, 표적 부위에서 또는 근방에서 적어도 하나의 히스티딘의 변형, 및/또는 표적 부위에서 또는 근방에서 전사의 변화를 포함할 수 있다.
(a) 세포로의 도입
상기 언급한 바와 같이, 상기 방법은 적어도 하나의 조작된 Cas9 시스템 및/또는 상기 시스템을 인코딩하는 핵산 (및 선택적인 공여체 폴리뉴클레오티드)을 진핵 세포에 도입하는 것을 포함한다. 적어도 하나의 시스템 및/또는 핵산/공여체 폴리뉴클레오티드는 다양한 수단에 의해 관심 세포 내부에 도입될 수 있다.
일부 구체예들에서, 상기 세포는 적절한 분자들 (즉, 단백질, DNA, 및/또는 RNA)로 형질감염 될 수 있다. 적합한 형질감염 방법들에는 뉴클레오펙션 (또는 전기천공), 칼슘 포스페이트-매개된 형질감염, 양이온 폴리머 형질감염 (예컨대, DEAE-덱스트란 또는 폴리에틸렌이민), 바이러스 형질도입, 비로좀 형질감염, 비리온 형질감염, 리포좀 형질감염, 양이온 리포좀 형질감염, 면역리포좀 형질감염, 비리포좀 지질 형질감염, 덴드리머 형질감염, 열 충격 형질감염, 자기주입법, 리포펙션, 유전자 총 전달, 임팔레펙션, 초음파천공법, 광학 형질감염, 및 특정상표 제제 (proprietary agent)-개선된 핵산의 흡수가 포함된다. 형질감염 방법들은 해당 분야에 잘 공지되어 있다 (예컨대, “Current Protocols in Molecular Biology” Ausubel 외., John Wiley & Sons, New York, 2003 or “Molecular Cloning: A Laboratory Manual” Sambrook & Russell, Cold Spring Harbor Press, Cold Spring Harbor, NY, 3rd edition, 2001 참고). 다른 구체예들에서, 상기 분자들은 미세주입에 의하여 세포 내부에 도입될 수 있다. 예를 들면, 상기 분자들은 관심 세포들의 세포질 또는 핵 내부에 주사될 수 있다. 세포 내부에 도입되는 각 분자의 양은 변화할 수 있으나, 해당 분야의 당업자는 적절한 양을 결정하는 수단을 잘 알고 있다.
다양한 분자들은 동시에 또는 순차적으로 세포 내부에 도입될 수 있다. 예를 들어, 조작된 Cas9 시스템 (또는 이의 인코딩 핵산)과 공여체 폴리뉴클레오티드는 동시에 도입 될 수 있다. 대안적으로, 하나를 먼저 도입한 다음 다른 하나를 차후에 세포 내에 도입할 수 있다.
일반적으로, 세포는 세포 성장 및/또는 유지에 적절한 조건하에서 유지된다. 적합한 세포 배양 조건들은 해당 분야에 널리 공지이며, 예를 들면, Santiago 외., Proc. Natl. Acad. Sci. USA, 2008, 105:5809-5814; Moehle 외. Proc. Natl. Acad. Sci. USA, 2007, 104:3055-3060; Urnov 외., Nature, 2005, 435:646-651; 및 Lombardo 외., Nat. Biotechnol., 2007, 25:1298-1306에 기재되어 있다. 해당 분야의 당업자들은 세포의 배양 방법들이 해당 분야에 공지이며 세포 유형에 따라 달라질 것임을 이해하고 있을 것이다. 모든 사례에서, 특정 세포 유형에 대한 가장 우수한 기술들을 결정하기 위해 관례적인 최적화가 사용될 수 있다.
(b) 선택적 공여체 폴리뉴클레오티드
조작된 Cas9 단백질이 뉴클레아제 또는 니카아제 활성을 포함하는 구체예들에서, 상기 방법은 최소한 하나의 공여체 폴리뉴클레오티드를 세포에 도입하는 단계를 추가로 포함할 수 있다. 공여체 폴리뉴클레오티드는 단일-가닥 또는 이중-가닥, 선형 또는 원형, 및/또는 RNA 또는 DNA 일 수 있다. 일부 구체예들에서, 공여체 폴리뉴클레오티드는 벡터, 예컨대, 플라스미드 벡터일 수 있다.
공여체 폴리뉴클레오티드는 최소한 하나의 공여체 서열을 포함한다. 일부 양상들에서, 공여체 폴리뉴클레오티드의 공여체 서열은 내인성 또는 고유 염색체 서열의 변형된 형태일 수 있다. 예를 들면, 공여체 서열은 조작된 Cas9 시스템에 의해 표적되는 서열의 또는 서열 근방의 염색체 서열의 일부분과 실질적으로 동일할 수 있지만, 이는 최소한 하나의 뉴클레오티드 변화를 포함한다. 그러므로, 고유 서열과 통합 또는 교환시, 표적된 염색체 위치의 서열은 최소한 하나의 뉴클레오티드 변화를 포함한다. 예를 들면, 이러한 변화는 하나 이상의 뉴클레오티드 삽입, 하나 이상의 뉴클레오티드 결실, 하나 이상의 뉴클레오티드 치환, 또는 이의 조합일 수 있다. 변형된 서열의 “유전자 교정” 통합의 결과로서, 세포는 표적된 염색체 서열로부터 변형된 유전자 생성물을 생성할 수 있다.
다른 양상들에서, 공여체 폴리뉴클레오티드의 공여체 서열은 외인성 서열일 수 있다. 본 명세서에서 사용되는, “외인성” 서열은 세포에 대해 고유적이지 않은 서열, 또는 그 고유 위치가 세포의 게놈에서 상이한 위치에 존재하는 서열을 지칭한다. 예를 들면, 외인성 서열은 단백질 코딩 서열을 포함할 수 있는데, 이는 외인성 프로모터 제어 서열에 작동적으로 연결될 수 있으므로, 게놈에 통합시, 세포는 통합된 서열에 의해 코드되는 단백질을 발현할 수 있다. 대안적으로, 외인성 서열은 염색체 서열에 통합될 수 있으며, 그리하여 그 발현은 내인성 프로모터 제어 서열에 의해 조절된다. 다른 반복들에서, 외인성 서열은 전사 제어 서열, 또 다른 발현 제어 서열, RNA 코딩 서열, 등일 수 있다. 상기 언급한 바와 같이, 외인성 서열의 염색체 서열로의 통합은 “녹인”이라 명명한다.
해당 분야의 당업자가 이해할 수 있는 바와 같이, 공여체 서열의 길이는 달라질 수 있으며 달라질 것이다. 예를 들면, 공여체 서열은 수개의 뉴클레오티드에서 수백개의 뉴클레오티드까지의 길이에서 수십만개 뉴클레오티드까지 변화할 수 있다.
전형적으로, 공여체 폴리뉴클레오티드의 공여체 서열은 상류 서열 및 하류 서열에 의해 연접되고, 이는 각각 조작된 Cas9 시스템에 의해 표적되는 서열의 상류 및 하류에 위치한 서열들과 실질적 서열 동일성을 가진다. 이들 서열 유사성으로 인해, 공여체 폴리뉴클레오티드의 상류 및 하류 서열들은 공여체 폴리뉴클레오티드와 표적되는 염색체 서열 간의 상동 재조합을 가능하게 하여 공여체 서열이 염색체 서열 내에 통합 (또는 염색체 서열과 교환) 될 수 있다.
본 명세서에서 사용되는 상류 서열은, 조작된 Cas9 시스템에 의해 표적되는 서열의 상류 염색체 서열과 실질적인 서열 동일성을 공유하는 핵산 서열을 지칭한다. 유사하게, 하류 서열은 조작된 Cas9 시스템에 의해 표적되는 서열의 하류 염색체 서열과 실질적인 서열 동일성을 공유하는 핵산 서열을 지칭한다. 본 명세서에서 사용되는, 어구 “실질적 서열 동일성”은 최소한 약 75% 서열 동일성을 가지는 서열들을 지칭한다. 그러므로, 공여체 폴리뉴클레오티드 내 상류 및 하류 서열들은 표적 서열에 대한 상류 또는 하류 서열과 약 75%, 76%, 77%, 78%, 79%, 80%, 81%, 82%, 83%, 84%, 85%, 86%, 87%, 88%, 89%, 90%, 91%, 92%, 93%, 94%, 95%, 96%, 97%, 98%, 또는 99% 서열 동일성을 가질 수 있다. 한 예시적 구체예에서, 공여체 폴리뉴클레오티드 내 상류 및 하류 서열들은 조작된 Cas9 시스템에 의해 표적되는 서열에 대한 상류 또는 하류 염색체 서열과 약 95% 또는 100% 서열 동일성을 가질 수 있다.
일부 구체예들에서, 상류 서열은 조작된 Cas9 시스템에 의해 표적되는 서열의 바로 상류에 위치한 염색체 서열과 실질적 서열 동일성을 공유한다. 다른 구체예들에서, 상류 서열은 표적 서열로부터 약 백 (100)개 뉴클레오티드 상류 이내에 위치한 염색체 서열과 실질적 서열 동일성을 공유한다. 그러므로, 예를 들면, 상류 서열은 표적 서열로부터 약 1 내지 약 20, 약 21 내지 약 40, 약 41 내지 약 60, 약 61 내지 약 80, 또는 약 81 내지 약 100 뉴클레오티드 상류에 위치한 염색체 서열과 실질적 서열 동일성을 공유할 수 있다. 일부 구체예들에서, 하류 서열은 조작된 Cas9 시스템에 의해 표적되는 서열의 바로 하류에 위치한 염색체 서열과 실질적 서열 동일성을 공유한다. 다른 구체예들에서, 하류 서열은 표적 서열로부터 약 백 (100)개 뉴클레오티드 하류 이내에 위치한 염색체 서열과 실질적 서열 동일성을 공유한다. 그러므로, 예를 들면, 하류 서열은 표적 서열로부터 약 1 내지 약 20, 약 21 내지 약 40, 약 41 내지 약 60, 약 61 내지 약 80, 또는 약 81 내지 약 100 뉴클레오티드 하류에 위치한 염색체 서열과 실질적 서열 동일성을 공유할 수 있다.
각 상류 또는 하류 서열은 약 20개 뉴클레오티드 내지 약 5000개 뉴클레오티드 길이 범위일 수 있다. 일부 구체예들에서, 상류 및 하류 서열들은 약 50, 100, 200, 300, 400, 500, 600, 700, 800, 900, 1000, 1100, 1200, 1300, 1400, 1500, 1600, 1700, 1800, 1900, 2000, 2100, 2200, 2300, 2400, 2500, 2600, 2800, 3000, 3200, 3400, 3600, 3800, 4000, 4200, 4400, 4600, 4800, 또는 5000개 뉴클레오티드를 포함할 수 있다. 특정 구체예들에서, 상류 및 하류 서열들은 약 50 내지 약 1500개 뉴클레오티드 길이 범위일 수 있다.
(c) 세포 유형
다양한 진핵 세포들이 본 발명에 개시된 방법에서 사용하기에 적합하다. 예를 들면, 세포는 인간 세포, 비-인간 포유동물 세포, 비-포유동물 척추동물 세포, 무척추동물 세포, 곤충 세포, 식물 세포, 효모 세포, 또는 단세포 진핵생물 기관일 수 있다. 일부 구체예들에서, 세포는 하나의 세포 배아 일 수도 있다. 예를 들면, 비-인간 포유동물 배아는 쥐, 햄스터, 설치류, 토끼, 고양이, 개, 양, 돼지, 소, 말, 및 영장류 배아를 포함한다. 또 다른 구체예들에서, 세포는 줄기 세포, 가령, 배아 줄기 세포, ES-유사 줄기 세포, 태아 줄기 세포, 성체 줄기 세포, 등일 수 있다. 한 구체예에서, 줄기 세포는 인간 배아 줄기 세포가 아니다. 더욱이, 줄기 세포들은 그 전문이 본 명세서에 포함되는 WO2003/046141, 또는 Chung 외. (Cell 줄기 세포, 2008, 2:113-117)에 개시된 기술들에 의해 제조되는 것들을 포함할 수 있다. 세포는 시험관내 (즉, 배양물 내), 생체외 (즉, 유기체로부터 분리된 조직 내) 또는 생체내 (즉, 유기체내) 존재할 수 있다. 예시적 구체예들에서, 세포는 포유동물 세포 또는 포유동물 세포주이다. 특정 구체예들에서, 세포는 인간 세포 또는 인간 세포주이다.
예로서, 일부 구체예들에서, 진핵 세포 또는 진핵 세포 집단은 T 세포, CD8+ T 세포, CD8+ 나이브 T 세포, 중추 기억 T 세포, 효과기 기억 T 세포, CD4+ T 세포, 줄기 세포 기억 T 세포, 보조 T 세포, 조절 T 세포, 세포독성 T 세포, 자연 살해 T 세포, 조혈 줄기 세포, 장기 조혈 줄기 세포, 단기 조혈 줄기 세포, 다분화능 전구 세포, 계통 제한 전구 세포, 림프 전구 세포, 췌장 전구 세포, 내분비 전구 세포, 외분비 전구 세포, 골수 전구 세포, 일반 골수 전구 세포, 적혈구 전구 세포, 거핵 세포 적혈구 전구 세포, 단핵구 전구 세포, 내분비 전구 세포, 외분비 세포, 섬유아세포, 간모세포, 근모세포, 대식세포, 섬 베타 세포, 심근세포, 혈액 세포, 관 세포, 포상 세포, 알파 세포, 베타 세포, 델타 세포, PP 세포, 담관 세포, 망막 세포, 광수용기 세포, 간상 세포, 원추 세포, 망막 색소 상피 세포, 섬유주 세포, 와우 세포, 외유모 세포, 내유모 세포, 폐 상피 세포, 기관지 상피 세포, 폐포 상피 세포, 폐 상피 전구 세포, 횡문근 세포, 심장 근육 세포, 근육 위성 세포, 근세포, 뉴런, 뉴런 줄기 세포, 중간엽 줄기 세포, 유도 만능 줄기 세포(iPS) 세포 , 배아줄기세포, 단핵구, 거핵구, 호중구, 호산구, 호염기구, 비만세포, 망상적혈구, B 세포, 예를 들어, 전구 B 세포, 예를 들어, Pre B 세포, Pro B 세포, 기억 B 세포, 혈장 B 세포, 위장 상피 세포, 담도 상피 세포, 췌관 상피 세포, 장 줄기 세포, 간세포, 간 성상 세포, 쿠퍼 세포, 조골 세포, 파골 세포, 지방세포(예를 들어, 갈색 지방세포 또는 백색 지방세포), 전지방세포, 췌장 반응성 전구체 세포, 췌장 섬 세포, 췌장 베타 세포, 췌장 알파 세포, 췌장 델타 세포, 췌장 외분비 세포, 슈반 세포 또는 희돌기아교세포, 또는 이러한 세포의 집단이다.
적합한 포유동물 세포 또는 세포주들의 비-제한적 예들에는 인간 배아 신장 세포 (HEK293, HEK293T); 인간 자궁경부 암종 세포 (HELA); 인간 폐 세포 (W138); 인간 간 세포 (Hep G2); 인간 U2-OS 골육종 세포, 인간 A549 세포, 인간 A-431 세포, 및 인간 K562 세포; 중국 햄스터 난소 (CHO) 세포, 새끼 햄스터 신장 (BHK) 세포; 생쥐 골수종 NS0 세포, 생쥐 배아 섬유모세포 3T3 세포 (NIH3T3), 생쥐 B 림프종 A20 세포; 생쥐 흑색종 B16 세포; 생쥐 근육모세포 C2C12 세포; 생쥐 골수종 SP2/0 세포; 생쥐 배아 중간엽 C3H-10T1/2 세포; 생쥐 암종 CT26 세포, 생쥐 전립선 DuCuP 세포; 생쥐 유방 EMT6 세포; 생쥐 간암 Hepa1c1c7 세포; 생쥐 골수종 J5582 세포; 생쥐 상피 MTD-1A 세포; 생쥐 심근 MyEnd 세포; 생쥐 신장 RenCa 세포; 생쥐 이자 RIN-5F 세포; 생쥐 흑색종 X64 세포; 생쥐 림프종 YAC-1 세포; 쥐 교모세포종 9L 세포; 쥐 B 림프종 RBL 세포; 쥐 신경모세포종 B35 세포; 쥐 간암 세포 (HTC); 버팔로 쥐 간 BRL 3A 세포; 개 신장 세포 (MDCK); 개 유선 (CMT) 세포; 쥐 골육종 D17 세포; 쥐 단핵구/대식세포 DH82 세포; 원숭이 신장 SV-40 형질전환된 섬유모세포 (COS7) 세포; 원숭이 신장 CVI-76 세포; 아프리카 녹색 원숭이 신장 (VERO-76) 세포가 포함된다. 포유동물 세포주의 보다 많은 목록은 미국 표준 균주 카탈로그 (ATCC, Manassas, VA)에서 찾을 수 있다.
(V)
응용
본 명세서에 개시된 조성물 및 방법들은 다양한 치료, 진단, 산업 및 연구 분야에서 사용될 수 있다. 일부 구체예들에서, 본 발명은 유전자 기능을 모형화 및/또는 연구하기 위해, 관심의 유전적 또는 후성 조건들을 연구하기 위해, 또는 다양한 질병 또는 장애들에 관여하는 생화학적 경로들을 연구하기 위해 세포, 동물, 또는 식물에서 임의의 관심 염색체 서열을 변형하기 위해 사용될 수 있다. 예를 들면, 질병 또는 장애들을 모형화하는 유전자삽입 유기체가 생성될 수 있으며, 여기서 질병 또는 장애와 연관된 하나 이상의 핵산 서열들의 발현은 변화되어 있다. 질병 모델은 유기체에 대한 돌연변이들의 효과를 연구하기 위해, 질병의 발병 및/또는 진행을 연구하기 위해, 질병에 대한 제약학적 활성 화합물의 효과를 연구하기 위해, 및/또는 잠재적인 유전자 치료 전략의 효능을 평가하기 위해 사용될 수 있다.
다른 구체예들에서, 상기 조성물 및 방법들은 효율적인 그리고 비용 효과적인 기능적 게놈 선별을 실시하기 위해 사용될 수 있으며, 특정 생물학적 과정에 관여하는 유전자들의 기능 그리고 유전자 발현에서 임의의 변형이 생물학적 과정에 어떻게 영향을 줄 수 있는지를 연구하기 위해, 또는 세포 표현형과 관련된 게놈 좌위의 포화 또는 딥 스캐닝 돌연변이생성을 실시하기 위해 사용될 수 있다. 예를 들면, 유전자 발현, 약물 내성, 및 질병의 역전에 필요한 기능적 요소들에 관한 중요한 최소 특징들 및 별개의 취약성들을 결정하기 위하여 포화 또는 딥 스캐닝 돌연변이생성이 사용될 수 있다.
추가 구체예들에서, 본 명세서에 개시된 조성물 및 방법들은 질병 또는 장애의 존재를 확인하기 위한 진단 테스트에 및/또는 치료 옵션 결정에 사용하기 위해 사용될 수 있다. 적합한 진단 테스트들의 예는 암 세포에서 특정 돌연변이들 (예컨대, EGFR, HER2, 등에서 특정 돌연변이)의 탐지, 특정 질병들과 관련된 특정 돌연변이들의 탐지 (예컨대, 트라이뉴클레오티드 반복, 낫 적혈구병과 연관된 β-글로빈의 돌연변이들, 특정 SNP, 등), 간염의 탐지, 바이러스 (예컨대, Zika)의 탐지, 등을 포함한다.
추가 구체예들에서, 본 명세서에 개시된 조성물 및 방법들은 특정 질병 또는 장애와 연관된 유전자 돌연변이들을 교정, 가령, 예컨대, 낫 적혈구병 또는 지중해빈혈과 연관된 글로빈 유전자 돌연변이들을 교정, 중증 복합성 면역 결핍증 (SCID)과 연관된 아데노신 탈아미노효소 유전자에서 돌연변이들을 교정, 헌팅턴 병의 질병-유발 유전자인 HTT의 발현을 감소, 또는 망막 색소변성의 치료를 위해 로돕신 유전자에서의 돌연변이들을 교정함에 사용될 수 있다. 이러한 변형들은 생체외 세포에서 이루어질 수 있다.
또 다른 구체예들에서, 본 명세서에 개시된 조성물 및 방법들은 환경적 스트레스에 대한 내성이 증가된 또는 개선된 형질들을 가지는 작물들을 생성하기 위해 사용될 수 있다. 본 발명은 또한 개선된 형질을 가진 농장 동물 또는 생산 동물을 생성하기 위해 사용될 수도 있다. 예를 들면, 돼지는 생의학 모델, 특히, 재생 의학 또는 이종이식에서의 모델로서 매력적이게 하는 많은 특징들을 가진다.
또 다른 구체예에서, 본원에 개시된 조성물 및 방법을 사용하여 살아있는 세포 또는 화학적으로 고정된 세포 (포르말린-고정 파라핀 포매 임상 샘플에 사용되는 포르말린 고정) 내의 염색체 동일성 및 위치를 결정할 수 있다. 예를 들어, 본원에 개시된 펩티드 서열을 통해 형광성 단백질에 연결된 CRIPSR 복합체는 단일 또는 다중 사본으로 유전자 좌위에 표적화될 수 있고, 이러한 복합체는 염색체 좌위 사본 수 및/또는 위치를 결정하기 위해 현미경으로 탐지될 수 있다. 추적을 위한 예시적인 유전자 좌위는 중심체 영역, 텔로머 영역, 또는 단일의 동일한 CRISPR 복합체의 다중 사본이 결합할 수 있는 게놈의 기타 반복 영역을 포함할 수 있다.
정의
달리 정의되지 않는 한, 본 명세서에서 사용되는 모든 기술 및 과학 용어는 본 발명이 속하는 해당 분야의 기술자에 의해 일반적으로 이해되는 의미를 가진다. 다음 참고문헌들은 본 발명에서 사용되는 많은 용어들의 일반적인 정의를 당업자에게 제공한다: Singleton 등, Dictionary of Microbiology and Molecular Biology (2nd Ed. 1994); The Cambridge Dictionary of Science and Technology (Walker ed., 1988); The Glossary of Genetics, 5th Ed., R. Rieger 등 (eds.), Springer Verlag (1991); 및 Hale & Marham, The Harper Collins Dictionary of Biology (1991). 본 출원에서 사용되는, 하기 용어들은 달리 특정한 언급이 없는 한 하기 의미들을 가진다.
본 발명 또는 바람직한 구체예(들)의 구성요소들을 소개할 때, 관사 “하나 (a, an)", “그것” 및 “상기"는 하나 이상의 구성요소들이 존재함을 의미하는 것이다. 용어 “포함하는", “비롯한” 및 “가지는"은 포함적인 의미이며 나열된 구성요소들 이외에 추가 구성요소들이 존재할 수 있음을 의미한다.
수치값과 관련하여 사용될 때 용어 “약” x는, 예를 들면 x ± 5%를 의미한다.
본 명세서에서 사용되는 용어 “상보적인” 또는 “상보성”은 특정 수소 결합을 통한 염기 페어링에 의한 이중-가닥 핵산들의 결합을 지칭한다. 염기 페어링은 표준 왓슨-크릭 염기 페어링일 수 있다 (예컨대, 5'-A G T C-3'은 상보적 서열 3'-T C A G-5'과 쌍을 이룬다). 염기 페어링은 또한 후그스틴 또는 역 후그스틴 수소 결합 일 수 있다. 상보성은 일반적으로 이중나선 영역에 대해 측정되므로, 예를 들면, 오버행은 제외된다. 이중나선 영역의 두 개 가닥들 간의 상보성은 부분적일 수 있으며 염기들 중 일부만 (예컨대, 70%) 상보적인 경우 백분율로 표현된다 (예컨대, 70%). 상보적이지 않은 염기들은 “미스매치”된다. 상보성은 또한 이중나선 영역의 모든 염기들이 상보적인 경우 완전할 수도 있다 (즉, 100%).
본 명세서에서 사용되는 용어 “CRISPR/Cas 시스템” 또는 "Cas9 시스템"은 Cas9 단백질 (즉, 뉴클레아제, 니카아제, 또는 촉매적 사멸 단백질) 및 가이드 RNA를 포함하는 복합체를 지칭한다.
본 명세서에서 사용되는 용어 “내인성 서열”은 세포에 대해 고유한 염색체 서열을 지칭한다.
본 명세서에서 사용되는 용어 “외인성”은 세포에 대해 고유하지 않은 서열 또는 세포의 게놈에서 그 고유 위치가 상이한 염색체 위치에 존재하는 염색체 서열을 지칭한다.
본 명세서에서 사용되는 “유전자”는 유전자 생성물을 인코딩하는 DNA 영역 (엑손과 인트론 포함), 뿐만 아니라 조절 서열들이 코딩 및/또는 전사된 서열들에 인접하는지와 관계없이 유전자 생성물의 생성을 조절하는 모든 DNA 영역을 지칭한다. 따라서, 유전자는 프로모터 서열, 터미네이터, 번역 조절 서열, 가령, 리보솜 결합 부위 및 내부 리보솜 진입 부위, 인핸서, 슬라이서, 절연체, 경계 요소, 복제 원점, 기질 부착 부위 및 좌위 제어 영역을 포함하나 이에 제한되는 것은 아니다.
용어 “이종”은 관심 세포에 대해 내인성이 아닌 또는 고유하지 않은 엔터티를 지칭한다. 예를 들면, 이종 단백질은 외인성 출처, 가령, 외인적으로 도입된 핵산 서열로부터 유래한 또는 본래 유래하였던 단백질을 지칭한다. 일부 사례들에서, 이종 단백질은 관심 세포에 의하여 일반적으로 생성되지 않는다.
용어 “니카아제”는 이중-가닥 핵산 서열 중 한 가닥을 절단하는 (즉, 이중-가닥 서열을 절단시키는) 효소를 지칭한다. 예를 들면, 이중 가닥 절단 활성을 가지는 뉴클레아제는 니카아제로서 기능하여 이중-가닥 서열 중 한 가닥만을 절단하도록 하기 위한 돌연변이 및/또는 결실에 의해 변형될 수 있다.
본 명세서에서 사용되는 용어 “뉴클레아제”는 이중-가닥 핵산 서열 두 가닥 모두를 절단하는 효소를 지칭한다.
용어 “핵산” 및 “폴리뉴클레오티드”는 선형 또는 원형 입체형태의, 그리고 단일- 또는 이중-가닥 형태의 데옥시리보뉴클레오티드 또는 리보뉴클레오티드 폴리머를 지칭한다. 본 출원의 목적에서, 이들 용어들은 폴리머의 길이에 대한 제한으로 해석되어서는 안된다. 이 용어들은 자연 뉴클레오티드, 뿐만 아니라 염기, 당 및/또는 포스페이트 모이어티 (예컨대, 포스포로티오에이트 골격)에서 변형되어 있는 뉴클레오티드의 공지된 유사체를 포함할 수 있다. 일반적으로, 특정 뉴클레오티드의 유사체는 동일한 염기-쌍 특이성을 가진다; 즉, A의 유사체는 T와 염기-쌍을 이루게 될 것이다.
용어 "뉴클레오티드”는 데옥시리보뉴클레오티드 또는 리보뉴클레오티드를 지칭한다. 뉴클레오티드는 표준 뉴클레오티드 (즉, 아데노신, 구아노신, 시티딘, 티미딘, 및 우리딘), 뉴클레오티드 이성질체, 또는 뉴클레오티드 유사체일 수 있다. 뉴클레오티드 유사체는 변형된 퓨린 또는 피리미딘 염기 또는 변형된 리보오스 모이어티를 가지는 뉴클레오티드를 지칭한다. 뉴클레오티드 유사체는 자연 발생 뉴클레오티드 (예컨대, 이노신, 수도우리딘, 등) 또는 비-자연 발생 뉴클레오티드 일 수 있다. 뉴클레오티드의 당 또는 염기 모이어티에 대한 변형의 비-제한적 예들에는 아세틸 그룹, 아미노 그룹, 카르복실 그룹, 카르복시메틸 그룹, 하이드록실 그룹, 메틸 그룹, 포스포릴 그룹, 및 싸이올 그룹의 부가 (또는 제거), 뿐만 아니라 염기의 탄소 및 질소 원자들의 다른 원자들로의 치환 (예컨대, 7-데아자 퓨린)이 포함된다. 뉴클레오티드 유사체에는 또한 다이데옥시 뉴클레오티드, 2'-O-메틸 뉴클레오티드, 잠금 핵산 (LNA), 펩티드 핵산 (PNA), 및 모르포린이 포함된다.
용어 “폴리펩티드” 및 “단백질”은 호환적으로 사용되어 아미노산 잔기들의 폴리머를 지칭한다.
용어 “표적 서열”, “표적 염색체 서열” 및 “표적 부위”는 호환적으로 사용되어 조작된 Cas9 시스템이 표적되는 염색체 DNA 내 특정 서열, 및 조작된 Cas9 시스템이 DNA 또는 DNA와 관련된 단백질(들)을 변형시키는 부위를 지칭한다.
핵산 및 아미노산 서열 동일성을 결정하는 기술들은 해당 분야에 공지되어 있다. 전형적으로, 이러한 기술들은 유전자에 대한 mRNA의 뉴클레오티드 서열을 결정하는 단계 및/또는 이에 의해 인코딩되는 아미노산 서열을 결정하는 단계, 및 이들 서열들을 제 2 뉴클레오티드 또는 아미노산 서열과 비교하는 단계를 포함한다. 또한 게놈 서열들을 결정하고 이러한 방식으로 비교할 수 있다. 일반적으로, 동일성은 두 개의 폴리뉴클레오티드 또는 폴리펩티드 서열 각각의 정확한 뉴클레오티드-대-뉴클레오티드 또는 아미노산-대-아미노산 상응성을 지칭한다. 둘 이상의 서열들 (폴리뉴클레오티드 또는 아미노산)은 그 퍼센트 동일성을 결정함으로써 비교될 수 있다. 2개 서열들, 핵산 또는 아미노산 서열들의 동일성 백분율은, 2개의 정렬된 서열들 간의 일치 정합수(exact matches)를 더 짧은 서열들의 길이로 나누고 100을 곱한 것이다. 핵산 서열들에 대한 대략적인 정렬은 Smith 및 Waterman의 국소적 상동성 알고리즘, Advances in Applied Mathematics 2:482-489 (1981)에 의해 제공된다. 이 알고리즘은 미국 Washington, D.C.의 National Biomedical Research Foundation이 펴낸 Dayhoff, Atlas of Atlas of Protein Sequences and Structure, M. O. Dayhoff ed., 5 suppl. 3:353-358에 의해 개발되고, Gribskov, Nucl. Acids Res. 14(6):6745-6763 (1986)에 의해 정규화된 점수 행렬을 사용함으로써 아미노산 서열들에 적용될 수 있다. 서열의 동일성 백분율을 결정하기 위한 이러한 알고리즘의 예시적인 실행은 “BestFit” 유틸리티 응용 프로그램에서 Genetics Computer Group (Madison, Wis.)에 의해 제공된다. 서열들 간의 동일성 또는 유사성 백분율을 계산하는 그 외 적합한 프로그램들은 일반적으로 해당 기술분야에 공지이며, 예를 들어, 또다른 정렬 프로그램은 기본 매개변수와 함께 사용되는 BLAST가 있다. 예를 들어, BLASTN 및 BLASTP는 다음 기본 매개변수를 사용하여 이용될 수 있다: 유전 부호=표준; 필터=없음; 가닥=모두; 컷오프=60; 기대값=10; 매트릭스=BLOSUM62; 디스크립션=50 서열들; 분류=HIGH SCORE; 데이터베이스=비다중(non-redundant), GenBank+EMBL+DDBJ+PDB+ GenBank CDS translations+Swiss protein+Spupdate+PIR. 이들 프로그램들의 상세내용은 GenBank 웹사이트에서 찾을 수 있다.
본 발명의 범위에서 벗어나지 않고 상기 세포 및 방법들에서 다양한 변화가 이루어질 수 있으므로, 상기 상세한 설명 및 하기 제공되는 실시예에 포함된 모든 주제들은 설명적으로 해석되어야 하며 제한적 의미로 해석되어서는 안된다.
실시예
다음 실시예들은 본 발명의 특정 양상들을 설명한다.
실시예 1: GFP-SpCas9 및 RFP-SpCas9 융합 단백질을 사용한 인간 세포 유전자 편집
인간 K562 세포 (0.35 x 106)는 60 pmol의 SpCas9, GFP-SpCas9 또는 RFP-SpCas9 재조합 단백질 및 가이드 서열 5'-GCUCCCAUCACAUCAACCGG-3'을 가지는 인간 EMX1 좌위를 표적하는 180 pmol의 시험관내 전사된 단일 가이드 RNA(sgRNA)로 형질감염되었다. Nucleofection Solution V 및 Amaxa 기기를 사용하여 형질감염을 수행했다. 세포는 유전자 편집 분석을 위해 수확하기 전 3일 동안 37oC 및 5% CO2에서 유지되었다. QuickExtract DNA 추출 용액을 사용하여 게놈 DNA를 준비했다. 표적 EMX1 영역은 표적 특이적 서열과 차세대 시퀀싱(NGS) 어댑터로 구성된 프라이머를 사용하여 PCR 증폭되었다. 정방향 프라이머는 5'- TCGTCGGCAGCGTCAGATGTGTATAAGAGACAGNNNNNNAGTCTTCCCATCAGGCTCTCA-3' (서열 번호: 46)이고 역방향 프라이머는 GTCTCGTGGGCTCGGAGATGTGTATAAGAGACAGNNNNNNAGAGTCCAGCTTGGGCC-3' (서열 번호: 47)이고, 이때 표적 특이적 서열은 밑줄표시 되고 N은 A, T, G 또는 C를 나타낸다. PCR 앰플리콘들을 Illumina MiSeq를 사용하는 NGS로 분석하여, 각 Cas9 단백질의 편집 효율을 결정하였다. 도 1에 도시된 결과는 GFP-SpCas9 및 RFP-SpCas9 융합 단백질 각각이 SpCas9 단백질에 의한 수준과 유사한 편집 활성을 유지함을 보여준다.
표 1은 조작된 Cas9/NLS 단백질의 인간 코돈 최적화된 DNA 및 단백질 서열을 제시하며, 여기서 NLS 서열은 볼드체 텍스트로 제시되고 마커 단백질과 Cas9 사이의 링커는 밑줄친 텍스트로 제시된다.
세 가지 단백질을 제조하는 데 사용되는 인간 코돈 최적화 DNA 서열은 다음과 같다:
실시예 2:
시판 제품들과 편집 효율성 비교
두 가지 시판 GFP-SpCas9 융합 단백질 제품들, GenCrispr NLS-Cas9-EGFP 뉴클레아제 및 ArciTect Cas9-eGFP 뉴클레아제를 각각 GenScript (Piscataway, NJ) 및 Stemcell Technologies (Vancouver, Canada)사로부터 구입하였다. 인간 U2OS 세포 (0.2 x 106) 및 HEK293 세포 (0.3 x 106)를, 각각 인간 EMX1, HEKSite4, VEGFA3, HPRT 유전자좌를 표적으로 하는 4개의 화학적으로 합성된 sgRNA 150 pmol와 조합하여, 50 pmol의 GenCrispr NLS-Cas9-EGFP 뉴클레아제 또는 ArciTect Cas9-eGFP 뉴클레아제 또는 본 발명의 GFP-SpCas9 단백질로 형질감염시켰다. 가이드 서열들은 다음과 같다: 5'-GAGUCCGAGCAGAAGAAGAA-3' (EMX1) (서열 번호: 51), 5'-GGCACUGCGGCUGGAGGUGG-3' (HEKSite4) (서열 번호: 52), 5'GGUGAGUGAGUGUGUGCGUG-3' (VEGFA3), 및 5'-GGUCACUUUUAACACACCCA-3' (HPRT) (서열 번호: 53). Nucleofection Solution V 및 Amaxa 기기를 사용하여 형질감염을 수행했다. 세포는 유전자 편집 분석을 위해 수확하기 전 3일 동안 37oC 및 5% CO2에서 유지되었다. QuickExtract DNA 추출 용액을 사용하여 게놈 DNA를 준비했다. 각각의 표적된 게놈 영역은 표적 특이적 서열과 차세대 시퀀싱(NGS) 어댑터로 구성된 한 쌍의 프라이머를 사용하여 PCR 증폭되었다. 프라이머들을 다음 표에 열거한다:
PCR 앰플리콘들을 Illumina MiSeq를 사용하는 NGS로 분석하여, 각 Cas9 단백질의 편집 효율을 결정하였다. 도 2A 및 도 2B의 결과는 본 발명의 GFP-SpCas9 단백질에 의한 편집 효율이 모든 표적에서 시판 단백질의 편집 효율보다 몇 배 더 높았음을 보여준다.
SEQUENCE LISTING
<110> SIGMA-ALDRICH CO. LLC
<120> CRISPR/CAS FUSION PROTEINS AND SYSTEMS
<130> P19-027 WO-PCT
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<150> 62/806,708
<151> 2019-02-15
<160> 66
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<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 14
Arg Glu Lys Lys Lys Phe Leu Lys Arg Arg
1 5 10
<210> 15
<211> 20
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 15
Lys Arg Lys Gly Asp Glu Val Asp Gly Val Asp Glu Val Ala Lys Lys
1 5 10 15
Lys Ser Lys Lys
20
<210> 16
<211> 17
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 16
Arg Lys Cys Leu Gln Ala Gly Met Asn Leu Glu Ala Arg Lys Thr Lys
1 5 10 15
Lys
<210> 17
<211> 38
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 17
Asn Gln Ser Ser Asn Phe Gly Pro Met Lys Gly Gly Asn Phe Gly Gly
1 5 10 15
Arg Ser Ser Gly Pro Tyr Gly Gly Gly Gly Gln Tyr Phe Ala Lys Pro
20 25 30
Arg Asn Gln Gly Gly Tyr
35
<210> 18
<211> 42
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 18
Arg Met Arg Ile Glx Phe Lys Asn Lys Gly Lys Asp Thr Ala Glu Leu
1 5 10 15
Arg Arg Arg Arg Val Glu Val Ser Val Glu Leu Arg Lys Ala Lys Lys
20 25 30
Asp Glu Gln Ile Leu Lys Arg Arg Asn Val
35 40
<210> 19
<211> 239
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 19
Met Val Ser Lys Gly Glu Glu Leu Phe Thr Gly Val Val Pro Ile Leu
1 5 10 15
Val Glu Leu Asp Gly Asp Val Asn Gly His Lys Phe Ser Val Ser Gly
20 25 30
Glu Gly Glu Gly Asp Ala Thr Tyr Gly Lys Leu Thr Leu Lys Phe Ile
35 40 45
Cys Thr Thr Gly Lys Leu Pro Val Pro Trp Pro Thr Leu Val Thr Thr
50 55 60
Leu Thr Tyr Gly Val Gln Cys Phe Ser Arg Tyr Pro Asp His Met Lys
65 70 75 80
Gln His Asp Phe Phe Lys Ser Ala Met Pro Glu Gly Tyr Val Gln Glu
85 90 95
Arg Thr Ile Phe Phe Lys Asp Asp Gly Asn Tyr Lys Thr Arg Ala Glu
100 105 110
Val Lys Phe Glu Gly Asp Thr Leu Val Asn Arg Ile Glu Leu Lys Gly
115 120 125
Ile Asp Phe Lys Glu Asp Gly Asn Ile Leu Gly His Lys Leu Glu Tyr
130 135 140
Asn Tyr Asn Ser His Asn Val Tyr Ile Met Ala Asp Lys Gln Lys Asn
145 150 155 160
Gly Ile Lys Val Asn Phe Lys Ile Arg His Asn Ile Glu Asp Gly Ser
165 170 175
Val Gln Leu Ala Asp His Tyr Gln Gln Asn Thr Pro Ile Gly Asp Gly
180 185 190
Pro Val Leu Leu Pro Asp Asn His Tyr Leu Ser Thr Gln Ser Lys Leu
195 200 205
Ser Lys Asp Pro Asn Glu Lys Arg Asp His Met Val Leu Leu Glu Phe
210 215 220
Val Thr Ala Ala Gly Ile Thr Leu Gly Met Asp Glu Leu Tyr Lys
225 230 235
<210> 20
<211> 232
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 20
Met Val Ser Lys Gly Glu Ala Val Ile Lys Glu Phe Met Arg Phe Lys
1 5 10 15
Val His Met Glu Gly Ser Met Asn Gly His Glu Phe Glu Ile Glu Gly
20 25 30
Glu Gly Glu Gly Arg Pro Tyr Glu Gly Thr Gln Thr Ala Lys Leu Lys
35 40 45
Val Thr Lys Gly Gly Pro Leu Pro Phe Ser Trp Asp Ile Leu Ser Pro
50 55 60
Gln Phe Met Tyr Gly Ser Arg Ala Phe Thr Lys His Pro Ala Asp Ile
65 70 75 80
Pro Asp Tyr Tyr Lys Gln Ser Phe Pro Glu Gly Phe Lys Trp Glu Arg
85 90 95
Val Met Asn Phe Glu Asp Gly Gly Ala Val Thr Val Thr Gln Asp Thr
100 105 110
Ser Leu Glu Asp Gly Thr Leu Ile Tyr Lys Val Lys Leu Arg Gly Thr
115 120 125
Asn Phe Pro Pro Asp Gly Pro Val Met Gln Lys Lys Thr Met Gly Trp
130 135 140
Glu Ala Ser Thr Glu Arg Leu Tyr Pro Glu Asp Gly Val Leu Lys Gly
145 150 155 160
Asp Ile Lys Met Ala Leu Arg Leu Lys Asp Gly Gly Arg Tyr Leu Ala
165 170 175
Asp Phe Lys Thr Thr Tyr Lys Ala Lys Lys Pro Val Gln Met Pro Gly
180 185 190
Ala Tyr Asn Val Asp Arg Lys Leu Asp Ile Thr Ser His Asn Glu Asp
195 200 205
Tyr Thr Val Val Glu Gln Tyr Glu Arg Ser Glu Gly Arg His Ser Thr
210 215 220
Gly Gly Met Asp Glu Leu Tyr Lys
225 230
<210> 21
<211> 8
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 21
Asp Tyr Lys Asp Asp Asp Asp Lys
1 5
<210> 22
<211> 6
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
6xHis tag"
<400> 22
His His His His His His
1 5
<210> 23
<211> 20
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 23
Gly Arg Lys Lys Arg Arg Gln Arg Arg Arg Pro Pro Gln Pro Lys Lys
1 5 10 15
Lys Arg Lys Val
20
<210> 24
<211> 19
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 24
Pro Leu Ser Ser Ile Phe Ser Arg Ile Gly Asp Pro Pro Lys Lys Lys
1 5 10 15
Arg Lys Val
<210> 25
<211> 24
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 25
Gly Ala Leu Phe Leu Gly Trp Leu Gly Ala Ala Gly Ser Thr Met Gly
1 5 10 15
Ala Pro Lys Lys Lys Arg Lys Val
20
<210> 26
<211> 27
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 26
Gly Ala Leu Phe Leu Gly Phe Leu Gly Ala Ala Gly Ser Thr Met Gly
1 5 10 15
Ala Trp Ser Gln Pro Lys Lys Lys Arg Lys Val
20 25
<210> 27
<211> 21
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 27
Lys Glu Thr Trp Trp Glu Thr Trp Trp Thr Glu Trp Ser Gln Pro Lys
1 5 10 15
Lys Lys Arg Lys Val
20
<210> 28
<211> 11
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 28
Tyr Ala Arg Ala Ala Ala Arg Gln Ala Arg Ala
1 5 10
<210> 29
<211> 11
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 29
Thr His Arg Leu Pro Arg Arg Arg Arg Arg Arg
1 5 10
<210> 30
<211> 11
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 30
Gly Gly Arg Arg Ala Arg Arg Arg Arg Arg Arg
1 5 10
<210> 31
<211> 12
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 31
Arg Arg Gln Arg Arg Thr Ser Lys Leu Met Lys Arg
1 5 10
<210> 32
<211> 27
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 32
Gly Trp Thr Leu Asn Ser Ala Gly Tyr Leu Leu Gly Lys Ile Asn Leu
1 5 10 15
Lys Ala Leu Ala Ala Leu Ala Lys Lys Ile Leu
20 25
<210> 33
<211> 33
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 33
Lys Ala Leu Ala Trp Glu Ala Lys Leu Ala Lys Ala Leu Ala Lys Ala
1 5 10 15
Leu Ala Lys His Leu Ala Lys Ala Leu Ala Lys Ala Leu Lys Cys Glu
20 25 30
Ala
<210> 34
<211> 16
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 34
Arg Gln Ile Lys Ile Trp Phe Gln Asn Arg Arg Met Lys Trp Lys Lys
1 5 10 15
<210> 35
<211> 46
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 35
Ala Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys
1 5 10 15
Glu Ala Ala Ala Lys Ala Leu Glu Ala Glu Ala Ala Ala Lys Glu Ala
20 25 30
Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Ala
35 40 45
<210> 36
<211> 32
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 36
Ser Gly Gly Ser Ser Gly Gly Ser Ser Gly Ser Glu Thr Pro Gly Thr
1 5 10 15
Ser Glu Ser Ala Thr Pro Glu Ser Ser Gly Gly Ser Ser Gly Gly Ser
20 25 30
<210> 37
<211> 6
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 37
Leu Glu Gly Gly Gly Ser
1 5
<210> 38
<211> 4
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 38
Thr Gly Ser Gly
1
<210> 39
<211> 8
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 39
Gly Gly Ser Gly Gly Gly Ser Gly
1 5
<210> 40
<211> 20
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<220>
<221> SITE
<222> (1)..(20)
<223> /note="This sequence may encompass 1-4 'Gly Gly Gly Gly Ser'
repeating units"
<400> 40
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Gly
1 5 10 15
Gly Gly Gly Ser
20
<210> 41
<211> 8
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<220>
<221> SITE
<222> (1)..(8)
<223> /note="This sequence may encompass 6-8 residues"
<400> 41
Gly Gly Gly Gly Gly Gly Gly Gly
1 5
<210> 42
<211> 20
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<220>
<221> SITE
<222> (1)..(20)
<223> /note="This sequence may encompass 1-4 'Glu Ala Ala Ala Lys'
repeating units"
<400> 42
Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu
1 5 10 15
Ala Ala Ala Lys
20
<210> 43
<211> 27
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<220>
<221> SITE
<222> (2)..(26)
<223> /note="This region may encompass 2-5 'Glu Ala Ala Ala Lys'
repeating units"
<400> 43
Ala Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys
1 5 10 15
Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Ala
20 25
<210> 44
<211> 5
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<400> 44
Pro Ala Pro Ala Pro
1 5
<210> 45
<211> 16
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
peptide"
<220>
<221> SITE
<222> (1)..(16)
<223> /note="This sequence may encompass 6-8 'Ala Pro'
repeating units"
<400> 45
Ala Pro Ala Pro Ala Pro Ala Pro Ala Pro Ala Pro Ala Pro Ala Pro
1 5 10 15
<210> 46
<211> 60
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (34)..(39)
<223> a, t, g or c
<400> 46
tcgtcggcag cgtcagatgt gtataagaga cagnnnnnna gtcttcccat caggctctca 60
<210> 47
<211> 57
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (35)..(40)
<223> a, t, g or c
<400> 47
gtctcgtggg ctcggagatg tgtataagag acagnnnnnn agagtccagc ttgggcc 57
<210> 48
<211> 1692
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 48
Met Val Ser Lys Gly Glu Glu Leu Phe Thr Gly Val Val Pro Ile Leu
1 5 10 15
Val Glu Leu Asp Gly Asp Val Asn Gly His Lys Phe Ser Val Ser Gly
20 25 30
Glu Gly Glu Gly Asp Ala Thr Tyr Gly Lys Leu Thr Leu Lys Phe Ile
35 40 45
Cys Thr Thr Gly Lys Leu Pro Val Pro Trp Pro Thr Leu Val Thr Thr
50 55 60
Leu Thr Tyr Gly Val Gln Cys Phe Ser Arg Tyr Pro Asp His Met Lys
65 70 75 80
Gln His Asp Phe Phe Lys Ser Ala Met Pro Glu Gly Tyr Val Gln Glu
85 90 95
Arg Thr Ile Phe Phe Lys Asp Asp Gly Asn Tyr Lys Thr Arg Ala Glu
100 105 110
Val Lys Phe Glu Gly Asp Thr Leu Val Asn Arg Ile Glu Leu Lys Gly
115 120 125
Ile Asp Phe Lys Glu Asp Gly Asn Ile Leu Gly His Lys Leu Glu Tyr
130 135 140
Asn Tyr Asn Ser His Asn Val Tyr Ile Met Ala Asp Lys Gln Lys Asn
145 150 155 160
Gly Ile Lys Val Asn Phe Lys Ile Arg His Asn Ile Glu Asp Gly Ser
165 170 175
Val Gln Leu Ala Asp His Tyr Gln Gln Asn Thr Pro Ile Gly Asp Gly
180 185 190
Pro Val Leu Leu Pro Asp Asn His Tyr Leu Ser Thr Gln Ser Lys Leu
195 200 205
Ser Lys Asp Pro Asn Glu Lys Arg Asp His Met Val Leu Leu Glu Phe
210 215 220
Val Thr Ala Ala Gly Ile Thr Leu Gly Met Asp Glu Leu Tyr Lys Val
225 230 235 240
Asp Ala Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala
245 250 255
Lys Glu Ala Ala Ala Lys Ala Leu Glu Ala Glu Ala Ala Ala Lys Glu
260 265 270
Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Ala Pro
275 280 285
Ala Ala Lys Arg Val Lys Leu Asp Gly Gly Gly Gly Ser Thr Gly Met
290 295 300
Asp Lys Lys Tyr Ser Ile Gly Leu Asp Ile Gly Thr Asn Ser Val Gly
305 310 315 320
Trp Ala Val Ile Thr Asp Glu Tyr Lys Val Pro Ser Lys Lys Phe Lys
325 330 335
Val Leu Gly Asn Thr Asp Arg His Ser Ile Lys Lys Asn Leu Ile Gly
340 345 350
Ala Leu Leu Phe Asp Ser Gly Glu Thr Ala Glu Ala Thr Arg Leu Lys
355 360 365
Arg Thr Ala Arg Arg Arg Tyr Thr Arg Arg Lys Asn Arg Ile Cys Tyr
370 375 380
Leu Gln Glu Ile Phe Ser Asn Glu Met Ala Lys Val Asp Asp Ser Phe
385 390 395 400
Phe His Arg Leu Glu Glu Ser Phe Leu Val Glu Glu Asp Lys Lys His
405 410 415
Glu Arg His Pro Ile Phe Gly Asn Ile Val Asp Glu Val Ala Tyr His
420 425 430
Glu Lys Tyr Pro Thr Ile Tyr His Leu Arg Lys Lys Leu Val Asp Ser
435 440 445
Thr Asp Lys Ala Asp Leu Arg Leu Ile Tyr Leu Ala Leu Ala His Met
450 455 460
Ile Lys Phe Arg Gly His Phe Leu Ile Glu Gly Asp Leu Asn Pro Asp
465 470 475 480
Asn Ser Asp Val Asp Lys Leu Phe Ile Gln Leu Val Gln Thr Tyr Asn
485 490 495
Gln Leu Phe Glu Glu Asn Pro Ile Asn Ala Ser Gly Val Asp Ala Lys
500 505 510
Ala Ile Leu Ser Ala Arg Leu Ser Lys Ser Arg Arg Leu Glu Asn Leu
515 520 525
Ile Ala Gln Leu Pro Gly Glu Lys Lys Asn Gly Leu Phe Gly Asn Leu
530 535 540
Ile Ala Leu Ser Leu Gly Leu Thr Pro Asn Phe Lys Ser Asn Phe Asp
545 550 555 560
Leu Ala Glu Asp Ala Lys Leu Gln Leu Ser Lys Asp Thr Tyr Asp Asp
565 570 575
Asp Leu Asp Asn Leu Leu Ala Gln Ile Gly Asp Gln Tyr Ala Asp Leu
580 585 590
Phe Leu Ala Ala Lys Asn Leu Ser Asp Ala Ile Leu Leu Ser Asp Ile
595 600 605
Leu Arg Val Asn Thr Glu Ile Thr Lys Ala Pro Leu Ser Ala Ser Met
610 615 620
Ile Lys Arg Tyr Asp Glu His His Gln Asp Leu Thr Leu Leu Lys Ala
625 630 635 640
Leu Val Arg Gln Gln Leu Pro Glu Lys Tyr Lys Glu Ile Phe Phe Asp
645 650 655
Gln Ser Lys Asn Gly Tyr Ala Gly Tyr Ile Asp Gly Gly Ala Ser Gln
660 665 670
Glu Glu Phe Tyr Lys Phe Ile Lys Pro Ile Leu Glu Lys Met Asp Gly
675 680 685
Thr Glu Glu Leu Leu Val Lys Leu Asn Arg Glu Asp Leu Leu Arg Lys
690 695 700
Gln Arg Thr Phe Asp Asn Gly Ser Ile Pro His Gln Ile His Leu Gly
705 710 715 720
Glu Leu His Ala Ile Leu Arg Arg Gln Glu Asp Phe Tyr Pro Phe Leu
725 730 735
Lys Asp Asn Arg Glu Lys Ile Glu Lys Ile Leu Thr Phe Arg Ile Pro
740 745 750
Tyr Tyr Val Gly Pro Leu Ala Arg Gly Asn Ser Arg Phe Ala Trp Met
755 760 765
Thr Arg Lys Ser Glu Glu Thr Ile Thr Pro Trp Asn Phe Glu Glu Val
770 775 780
Val Asp Lys Gly Ala Ser Ala Gln Ser Phe Ile Glu Arg Met Thr Asn
785 790 795 800
Phe Asp Lys Asn Leu Pro Asn Glu Lys Val Leu Pro Lys His Ser Leu
805 810 815
Leu Tyr Glu Tyr Phe Thr Val Tyr Asn Glu Leu Thr Lys Val Lys Tyr
820 825 830
Val Thr Glu Gly Met Arg Lys Pro Ala Phe Leu Ser Gly Glu Gln Lys
835 840 845
Lys Ala Ile Val Asp Leu Leu Phe Lys Thr Asn Arg Lys Val Thr Val
850 855 860
Lys Gln Leu Lys Glu Asp Tyr Phe Lys Lys Ile Glu Cys Phe Asp Ser
865 870 875 880
Val Glu Ile Ser Gly Val Glu Asp Arg Phe Asn Ala Ser Leu Gly Thr
885 890 895
Tyr His Asp Leu Leu Lys Ile Ile Lys Asp Lys Asp Phe Leu Asp Asn
900 905 910
Glu Glu Asn Glu Asp Ile Leu Glu Asp Ile Val Leu Thr Leu Thr Leu
915 920 925
Phe Glu Asp Arg Glu Met Ile Glu Glu Arg Leu Lys Thr Tyr Ala His
930 935 940
Leu Phe Asp Asp Lys Val Met Lys Gln Leu Lys Arg Arg Arg Tyr Thr
945 950 955 960
Gly Trp Gly Arg Leu Ser Arg Lys Leu Ile Asn Gly Ile Arg Asp Lys
965 970 975
Gln Ser Gly Lys Thr Ile Leu Asp Phe Leu Lys Ser Asp Gly Phe Ala
980 985 990
Asn Arg Asn Phe Met Gln Leu Ile His Asp Asp Ser Leu Thr Phe Lys
995 1000 1005
Glu Asp Ile Gln Lys Ala Gln Val Ser Gly Gln Gly Asp Ser Leu
1010 1015 1020
His Glu His Ile Ala Asn Leu Ala Gly Ser Pro Ala Ile Lys Lys
1025 1030 1035
Gly Ile Leu Gln Thr Val Lys Val Val Asp Glu Leu Val Lys Val
1040 1045 1050
Met Gly Arg His Lys Pro Glu Asn Ile Val Ile Glu Met Ala Arg
1055 1060 1065
Glu Asn Gln Thr Thr Gln Lys Gly Gln Lys Asn Ser Arg Glu Arg
1070 1075 1080
Met Lys Arg Ile Glu Glu Gly Ile Lys Glu Leu Gly Ser Gln Ile
1085 1090 1095
Leu Lys Glu His Pro Val Glu Asn Thr Gln Leu Gln Asn Glu Lys
1100 1105 1110
Leu Tyr Leu Tyr Tyr Leu Gln Asn Gly Arg Asp Met Tyr Val Asp
1115 1120 1125
Gln Glu Leu Asp Ile Asn Arg Leu Ser Asp Tyr Asp Val Asp His
1130 1135 1140
Ile Val Pro Gln Ser Phe Leu Lys Asp Asp Ser Ile Asp Asn Lys
1145 1150 1155
Val Leu Thr Arg Ser Asp Lys Asn Arg Gly Lys Ser Asp Asn Val
1160 1165 1170
Pro Ser Glu Glu Val Val Lys Lys Met Lys Asn Tyr Trp Arg Gln
1175 1180 1185
Leu Leu Asn Ala Lys Leu Ile Thr Gln Arg Lys Phe Asp Asn Leu
1190 1195 1200
Thr Lys Ala Glu Arg Gly Gly Leu Ser Glu Leu Asp Lys Ala Gly
1205 1210 1215
Phe Ile Lys Arg Gln Leu Val Glu Thr Arg Gln Ile Thr Lys His
1220 1225 1230
Val Ala Gln Ile Leu Asp Ser Arg Met Asn Thr Lys Tyr Asp Glu
1235 1240 1245
Asn Asp Lys Leu Ile Arg Glu Val Lys Val Ile Thr Leu Lys Ser
1250 1255 1260
Lys Leu Val Ser Asp Phe Arg Lys Asp Phe Gln Phe Tyr Lys Val
1265 1270 1275
Arg Glu Ile Asn Asn Tyr His His Ala His Asp Ala Tyr Leu Asn
1280 1285 1290
Ala Val Val Gly Thr Ala Leu Ile Lys Lys Tyr Pro Lys Leu Glu
1295 1300 1305
Ser Glu Phe Val Tyr Gly Asp Tyr Lys Val Tyr Asp Val Arg Lys
1310 1315 1320
Met Ile Ala Lys Ser Glu Gln Glu Ile Gly Lys Ala Thr Ala Lys
1325 1330 1335
Tyr Phe Phe Tyr Ser Asn Ile Met Asn Phe Phe Lys Thr Glu Ile
1340 1345 1350
Thr Leu Ala Asn Gly Glu Ile Arg Lys Arg Pro Leu Ile Glu Thr
1355 1360 1365
Asn Gly Glu Thr Gly Glu Ile Val Trp Asp Lys Gly Arg Asp Phe
1370 1375 1380
Ala Thr Val Arg Lys Val Leu Ser Met Pro Gln Val Asn Ile Val
1385 1390 1395
Lys Lys Thr Glu Val Gln Thr Gly Gly Phe Ser Lys Glu Ser Ile
1400 1405 1410
Leu Pro Lys Arg Asn Ser Asp Lys Leu Ile Ala Arg Lys Lys Asp
1415 1420 1425
Trp Asp Pro Lys Lys Tyr Gly Gly Phe Asp Ser Pro Thr Val Ala
1430 1435 1440
Tyr Ser Val Leu Val Val Ala Lys Val Glu Lys Gly Lys Ser Lys
1445 1450 1455
Lys Leu Lys Ser Val Lys Glu Leu Leu Gly Ile Thr Ile Met Glu
1460 1465 1470
Arg Ser Ser Phe Glu Lys Asn Pro Ile Asp Phe Leu Glu Ala Lys
1475 1480 1485
Gly Tyr Lys Glu Val Lys Lys Asp Leu Ile Ile Lys Leu Pro Lys
1490 1495 1500
Tyr Ser Leu Phe Glu Leu Glu Asn Gly Arg Lys Arg Met Leu Ala
1505 1510 1515
Ser Ala Gly Glu Leu Gln Lys Gly Asn Glu Leu Ala Leu Pro Ser
1520 1525 1530
Lys Tyr Val Asn Phe Leu Tyr Leu Ala Ser His Tyr Glu Lys Leu
1535 1540 1545
Lys Gly Ser Pro Glu Asp Asn Glu Gln Lys Gln Leu Phe Val Glu
1550 1555 1560
Gln His Lys His Tyr Leu Asp Glu Ile Ile Glu Gln Ile Ser Glu
1565 1570 1575
Phe Ser Lys Arg Val Ile Leu Ala Asp Ala Asn Leu Asp Lys Val
1580 1585 1590
Leu Ser Ala Tyr Asn Lys His Arg Asp Lys Pro Ile Arg Glu Gln
1595 1600 1605
Ala Glu Asn Ile Ile His Leu Phe Thr Leu Thr Asn Leu Gly Ala
1610 1615 1620
Pro Ala Ala Phe Lys Tyr Phe Asp Thr Thr Ile Asp Arg Lys Arg
1625 1630 1635
Tyr Thr Ser Thr Lys Glu Val Leu Asp Ala Thr Leu Ile His Gln
1640 1645 1650
Ser Ile Thr Gly Leu Tyr Glu Thr Arg Ile Asp Leu Ser Gln Leu
1655 1660 1665
Gly Gly Asp Glu Phe Pro Lys Lys Lys Arg Lys Val Gly Gly Gly
1670 1675 1680
Gly Ser Pro Lys Lys Lys Arg Lys Val
1685 1690
<210> 49
<211> 1671
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 49
Met Val Ser Lys Gly Glu Ala Val Ile Lys Glu Phe Met Arg Phe Lys
1 5 10 15
Val His Met Glu Gly Ser Met Asn Gly His Glu Phe Glu Ile Glu Gly
20 25 30
Glu Gly Glu Gly Arg Pro Tyr Glu Gly Thr Gln Thr Ala Lys Leu Lys
35 40 45
Val Thr Lys Gly Gly Pro Leu Pro Phe Ser Trp Asp Ile Leu Ser Pro
50 55 60
Gln Phe Met Tyr Gly Ser Arg Ala Phe Thr Lys His Pro Ala Asp Ile
65 70 75 80
Pro Asp Tyr Tyr Lys Gln Ser Phe Pro Glu Gly Phe Lys Trp Glu Arg
85 90 95
Val Met Asn Phe Glu Asp Gly Gly Ala Val Thr Val Thr Gln Asp Thr
100 105 110
Ser Leu Glu Asp Gly Thr Leu Ile Tyr Lys Val Lys Leu Arg Gly Thr
115 120 125
Asn Phe Pro Pro Asp Gly Pro Val Met Gln Lys Lys Thr Met Gly Trp
130 135 140
Glu Ala Ser Thr Glu Arg Leu Tyr Pro Glu Asp Gly Val Leu Lys Gly
145 150 155 160
Asp Ile Lys Met Ala Leu Arg Leu Lys Asp Gly Gly Arg Tyr Leu Ala
165 170 175
Asp Phe Lys Thr Thr Tyr Lys Ala Lys Lys Pro Val Gln Met Pro Gly
180 185 190
Ala Tyr Asn Val Asp Arg Lys Leu Asp Ile Thr Ser His Asn Glu Asp
195 200 205
Tyr Thr Val Val Glu Gln Tyr Glu Arg Ser Glu Gly Arg His Ser Thr
210 215 220
Gly Gly Met Asp Glu Leu Tyr Lys Val Asp Ser Gly Gly Ser Ser Gly
225 230 235 240
Gly Ser Ser Gly Ser Glu Thr Pro Gly Thr Ser Glu Ser Ala Thr Pro
245 250 255
Glu Ser Ser Gly Gly Ser Ser Gly Gly Ser Pro Ala Ala Lys Arg Val
260 265 270
Lys Leu Asp Gly Gly Gly Gly Ser Thr Gly Met Asp Lys Lys Tyr Ser
275 280 285
Ile Gly Leu Asp Ile Gly Thr Asn Ser Val Gly Trp Ala Val Ile Thr
290 295 300
Asp Glu Tyr Lys Val Pro Ser Lys Lys Phe Lys Val Leu Gly Asn Thr
305 310 315 320
Asp Arg His Ser Ile Lys Lys Asn Leu Ile Gly Ala Leu Leu Phe Asp
325 330 335
Ser Gly Glu Thr Ala Glu Ala Thr Arg Leu Lys Arg Thr Ala Arg Arg
340 345 350
Arg Tyr Thr Arg Arg Lys Asn Arg Ile Cys Tyr Leu Gln Glu Ile Phe
355 360 365
Ser Asn Glu Met Ala Lys Val Asp Asp Ser Phe Phe His Arg Leu Glu
370 375 380
Glu Ser Phe Leu Val Glu Glu Asp Lys Lys His Glu Arg His Pro Ile
385 390 395 400
Phe Gly Asn Ile Val Asp Glu Val Ala Tyr His Glu Lys Tyr Pro Thr
405 410 415
Ile Tyr His Leu Arg Lys Lys Leu Val Asp Ser Thr Asp Lys Ala Asp
420 425 430
Leu Arg Leu Ile Tyr Leu Ala Leu Ala His Met Ile Lys Phe Arg Gly
435 440 445
His Phe Leu Ile Glu Gly Asp Leu Asn Pro Asp Asn Ser Asp Val Asp
450 455 460
Lys Leu Phe Ile Gln Leu Val Gln Thr Tyr Asn Gln Leu Phe Glu Glu
465 470 475 480
Asn Pro Ile Asn Ala Ser Gly Val Asp Ala Lys Ala Ile Leu Ser Ala
485 490 495
Arg Leu Ser Lys Ser Arg Arg Leu Glu Asn Leu Ile Ala Gln Leu Pro
500 505 510
Gly Glu Lys Lys Asn Gly Leu Phe Gly Asn Leu Ile Ala Leu Ser Leu
515 520 525
Gly Leu Thr Pro Asn Phe Lys Ser Asn Phe Asp Leu Ala Glu Asp Ala
530 535 540
Lys Leu Gln Leu Ser Lys Asp Thr Tyr Asp Asp Asp Leu Asp Asn Leu
545 550 555 560
Leu Ala Gln Ile Gly Asp Gln Tyr Ala Asp Leu Phe Leu Ala Ala Lys
565 570 575
Asn Leu Ser Asp Ala Ile Leu Leu Ser Asp Ile Leu Arg Val Asn Thr
580 585 590
Glu Ile Thr Lys Ala Pro Leu Ser Ala Ser Met Ile Lys Arg Tyr Asp
595 600 605
Glu His His Gln Asp Leu Thr Leu Leu Lys Ala Leu Val Arg Gln Gln
610 615 620
Leu Pro Glu Lys Tyr Lys Glu Ile Phe Phe Asp Gln Ser Lys Asn Gly
625 630 635 640
Tyr Ala Gly Tyr Ile Asp Gly Gly Ala Ser Gln Glu Glu Phe Tyr Lys
645 650 655
Phe Ile Lys Pro Ile Leu Glu Lys Met Asp Gly Thr Glu Glu Leu Leu
660 665 670
Val Lys Leu Asn Arg Glu Asp Leu Leu Arg Lys Gln Arg Thr Phe Asp
675 680 685
Asn Gly Ser Ile Pro His Gln Ile His Leu Gly Glu Leu His Ala Ile
690 695 700
Leu Arg Arg Gln Glu Asp Phe Tyr Pro Phe Leu Lys Asp Asn Arg Glu
705 710 715 720
Lys Ile Glu Lys Ile Leu Thr Phe Arg Ile Pro Tyr Tyr Val Gly Pro
725 730 735
Leu Ala Arg Gly Asn Ser Arg Phe Ala Trp Met Thr Arg Lys Ser Glu
740 745 750
Glu Thr Ile Thr Pro Trp Asn Phe Glu Glu Val Val Asp Lys Gly Ala
755 760 765
Ser Ala Gln Ser Phe Ile Glu Arg Met Thr Asn Phe Asp Lys Asn Leu
770 775 780
Pro Asn Glu Lys Val Leu Pro Lys His Ser Leu Leu Tyr Glu Tyr Phe
785 790 795 800
Thr Val Tyr Asn Glu Leu Thr Lys Val Lys Tyr Val Thr Glu Gly Met
805 810 815
Arg Lys Pro Ala Phe Leu Ser Gly Glu Gln Lys Lys Ala Ile Val Asp
820 825 830
Leu Leu Phe Lys Thr Asn Arg Lys Val Thr Val Lys Gln Leu Lys Glu
835 840 845
Asp Tyr Phe Lys Lys Ile Glu Cys Phe Asp Ser Val Glu Ile Ser Gly
850 855 860
Val Glu Asp Arg Phe Asn Ala Ser Leu Gly Thr Tyr His Asp Leu Leu
865 870 875 880
Lys Ile Ile Lys Asp Lys Asp Phe Leu Asp Asn Glu Glu Asn Glu Asp
885 890 895
Ile Leu Glu Asp Ile Val Leu Thr Leu Thr Leu Phe Glu Asp Arg Glu
900 905 910
Met Ile Glu Glu Arg Leu Lys Thr Tyr Ala His Leu Phe Asp Asp Lys
915 920 925
Val Met Lys Gln Leu Lys Arg Arg Arg Tyr Thr Gly Trp Gly Arg Leu
930 935 940
Ser Arg Lys Leu Ile Asn Gly Ile Arg Asp Lys Gln Ser Gly Lys Thr
945 950 955 960
Ile Leu Asp Phe Leu Lys Ser Asp Gly Phe Ala Asn Arg Asn Phe Met
965 970 975
Gln Leu Ile His Asp Asp Ser Leu Thr Phe Lys Glu Asp Ile Gln Lys
980 985 990
Ala Gln Val Ser Gly Gln Gly Asp Ser Leu His Glu His Ile Ala Asn
995 1000 1005
Leu Ala Gly Ser Pro Ala Ile Lys Lys Gly Ile Leu Gln Thr Val
1010 1015 1020
Lys Val Val Asp Glu Leu Val Lys Val Met Gly Arg His Lys Pro
1025 1030 1035
Glu Asn Ile Val Ile Glu Met Ala Arg Glu Asn Gln Thr Thr Gln
1040 1045 1050
Lys Gly Gln Lys Asn Ser Arg Glu Arg Met Lys Arg Ile Glu Glu
1055 1060 1065
Gly Ile Lys Glu Leu Gly Ser Gln Ile Leu Lys Glu His Pro Val
1070 1075 1080
Glu Asn Thr Gln Leu Gln Asn Glu Lys Leu Tyr Leu Tyr Tyr Leu
1085 1090 1095
Gln Asn Gly Arg Asp Met Tyr Val Asp Gln Glu Leu Asp Ile Asn
1100 1105 1110
Arg Leu Ser Asp Tyr Asp Val Asp His Ile Val Pro Gln Ser Phe
1115 1120 1125
Leu Lys Asp Asp Ser Ile Asp Asn Lys Val Leu Thr Arg Ser Asp
1130 1135 1140
Lys Asn Arg Gly Lys Ser Asp Asn Val Pro Ser Glu Glu Val Val
1145 1150 1155
Lys Lys Met Lys Asn Tyr Trp Arg Gln Leu Leu Asn Ala Lys Leu
1160 1165 1170
Ile Thr Gln Arg Lys Phe Asp Asn Leu Thr Lys Ala Glu Arg Gly
1175 1180 1185
Gly Leu Ser Glu Leu Asp Lys Ala Gly Phe Ile Lys Arg Gln Leu
1190 1195 1200
Val Glu Thr Arg Gln Ile Thr Lys His Val Ala Gln Ile Leu Asp
1205 1210 1215
Ser Arg Met Asn Thr Lys Tyr Asp Glu Asn Asp Lys Leu Ile Arg
1220 1225 1230
Glu Val Lys Val Ile Thr Leu Lys Ser Lys Leu Val Ser Asp Phe
1235 1240 1245
Arg Lys Asp Phe Gln Phe Tyr Lys Val Arg Glu Ile Asn Asn Tyr
1250 1255 1260
His His Ala His Asp Ala Tyr Leu Asn Ala Val Val Gly Thr Ala
1265 1270 1275
Leu Ile Lys Lys Tyr Pro Lys Leu Glu Ser Glu Phe Val Tyr Gly
1280 1285 1290
Asp Tyr Lys Val Tyr Asp Val Arg Lys Met Ile Ala Lys Ser Glu
1295 1300 1305
Gln Glu Ile Gly Lys Ala Thr Ala Lys Tyr Phe Phe Tyr Ser Asn
1310 1315 1320
Ile Met Asn Phe Phe Lys Thr Glu Ile Thr Leu Ala Asn Gly Glu
1325 1330 1335
Ile Arg Lys Arg Pro Leu Ile Glu Thr Asn Gly Glu Thr Gly Glu
1340 1345 1350
Ile Val Trp Asp Lys Gly Arg Asp Phe Ala Thr Val Arg Lys Val
1355 1360 1365
Leu Ser Met Pro Gln Val Asn Ile Val Lys Lys Thr Glu Val Gln
1370 1375 1380
Thr Gly Gly Phe Ser Lys Glu Ser Ile Leu Pro Lys Arg Asn Ser
1385 1390 1395
Asp Lys Leu Ile Ala Arg Lys Lys Asp Trp Asp Pro Lys Lys Tyr
1400 1405 1410
Gly Gly Phe Asp Ser Pro Thr Val Ala Tyr Ser Val Leu Val Val
1415 1420 1425
Ala Lys Val Glu Lys Gly Lys Ser Lys Lys Leu Lys Ser Val Lys
1430 1435 1440
Glu Leu Leu Gly Ile Thr Ile Met Glu Arg Ser Ser Phe Glu Lys
1445 1450 1455
Asn Pro Ile Asp Phe Leu Glu Ala Lys Gly Tyr Lys Glu Val Lys
1460 1465 1470
Lys Asp Leu Ile Ile Lys Leu Pro Lys Tyr Ser Leu Phe Glu Leu
1475 1480 1485
Glu Asn Gly Arg Lys Arg Met Leu Ala Ser Ala Gly Glu Leu Gln
1490 1495 1500
Lys Gly Asn Glu Leu Ala Leu Pro Ser Lys Tyr Val Asn Phe Leu
1505 1510 1515
Tyr Leu Ala Ser His Tyr Glu Lys Leu Lys Gly Ser Pro Glu Asp
1520 1525 1530
Asn Glu Gln Lys Gln Leu Phe Val Glu Gln His Lys His Tyr Leu
1535 1540 1545
Asp Glu Ile Ile Glu Gln Ile Ser Glu Phe Ser Lys Arg Val Ile
1550 1555 1560
Leu Ala Asp Ala Asn Leu Asp Lys Val Leu Ser Ala Tyr Asn Lys
1565 1570 1575
His Arg Asp Lys Pro Ile Arg Glu Gln Ala Glu Asn Ile Ile His
1580 1585 1590
Leu Phe Thr Leu Thr Asn Leu Gly Ala Pro Ala Ala Phe Lys Tyr
1595 1600 1605
Phe Asp Thr Thr Ile Asp Arg Lys Arg Tyr Thr Ser Thr Lys Glu
1610 1615 1620
Val Leu Asp Ala Thr Leu Ile His Gln Ser Ile Thr Gly Leu Tyr
1625 1630 1635
Glu Thr Arg Ile Asp Leu Ser Gln Leu Gly Gly Asp Glu Phe Pro
1640 1645 1650
Lys Lys Lys Arg Lys Val Gly Gly Gly Gly Ser Pro Lys Lys Lys
1655 1660 1665
Arg Lys Val
1670
<210> 50
<211> 1678
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 50
Met Val Ser Lys Gly Glu Glu Leu Phe Thr Gly Val Val Pro Ile Leu
1 5 10 15
Val Glu Leu Asp Gly Asp Val Asn Gly His Lys Phe Ser Val Ser Gly
20 25 30
Glu Gly Glu Gly Asp Ala Thr Tyr Gly Lys Leu Thr Leu Lys Phe Ile
35 40 45
Cys Thr Thr Gly Lys Leu Pro Val Pro Trp Pro Thr Leu Val Thr Thr
50 55 60
Leu Thr Tyr Gly Val Gln Cys Phe Ser Arg Tyr Pro Asp His Met Lys
65 70 75 80
Gln His Asp Phe Phe Lys Ser Ala Met Pro Glu Gly Tyr Val Gln Glu
85 90 95
Arg Thr Ile Phe Phe Lys Asp Asp Gly Asn Tyr Lys Thr Arg Ala Glu
100 105 110
Val Lys Phe Glu Gly Asp Thr Leu Val Asn Arg Ile Glu Leu Lys Gly
115 120 125
Ile Asp Phe Lys Glu Asp Gly Asn Ile Leu Gly His Lys Leu Glu Tyr
130 135 140
Asn Tyr Asn Ser His Asn Val Tyr Ile Met Ala Asp Lys Gln Lys Asn
145 150 155 160
Gly Ile Lys Val Asn Phe Lys Ile Arg His Asn Ile Glu Asp Gly Ser
165 170 175
Val Gln Leu Ala Asp His Tyr Gln Gln Asn Thr Pro Ile Gly Asp Gly
180 185 190
Pro Val Leu Leu Pro Asp Asn His Tyr Leu Ser Thr Gln Ser Lys Leu
195 200 205
Ser Lys Asp Pro Asn Glu Lys Arg Asp His Met Val Leu Leu Glu Phe
210 215 220
Val Thr Ala Ala Gly Ile Thr Leu Gly Met Asp Glu Leu Tyr Lys Val
225 230 235 240
Asp Ser Gly Gly Ser Ser Gly Gly Ser Ser Gly Ser Glu Thr Pro Gly
245 250 255
Thr Ser Glu Ser Ala Thr Pro Glu Ser Ser Gly Gly Ser Ser Gly Gly
260 265 270
Ser Pro Ala Ala Lys Arg Val Lys Leu Asp Gly Gly Gly Gly Ser Thr
275 280 285
Gly Met Asp Lys Lys Tyr Ser Ile Gly Leu Asp Ile Gly Thr Asn Ser
290 295 300
Val Gly Trp Ala Val Ile Thr Asp Glu Tyr Lys Val Pro Ser Lys Lys
305 310 315 320
Phe Lys Val Leu Gly Asn Thr Asp Arg His Ser Ile Lys Lys Asn Leu
325 330 335
Ile Gly Ala Leu Leu Phe Asp Ser Gly Glu Thr Ala Glu Ala Thr Arg
340 345 350
Leu Lys Arg Thr Ala Arg Arg Arg Tyr Thr Arg Arg Lys Asn Arg Ile
355 360 365
Cys Tyr Leu Gln Glu Ile Phe Ser Asn Glu Met Ala Lys Val Asp Asp
370 375 380
Ser Phe Phe His Arg Leu Glu Glu Ser Phe Leu Val Glu Glu Asp Lys
385 390 395 400
Lys His Glu Arg His Pro Ile Phe Gly Asn Ile Val Asp Glu Val Ala
405 410 415
Tyr His Glu Lys Tyr Pro Thr Ile Tyr His Leu Arg Lys Lys Leu Val
420 425 430
Asp Ser Thr Asp Lys Ala Asp Leu Arg Leu Ile Tyr Leu Ala Leu Ala
435 440 445
His Met Ile Lys Phe Arg Gly His Phe Leu Ile Glu Gly Asp Leu Asn
450 455 460
Pro Asp Asn Ser Asp Val Asp Lys Leu Phe Ile Gln Leu Val Gln Thr
465 470 475 480
Tyr Asn Gln Leu Phe Glu Glu Asn Pro Ile Asn Ala Ser Gly Val Asp
485 490 495
Ala Lys Ala Ile Leu Ser Ala Arg Leu Ser Lys Ser Arg Arg Leu Glu
500 505 510
Asn Leu Ile Ala Gln Leu Pro Gly Glu Lys Lys Asn Gly Leu Phe Gly
515 520 525
Asn Leu Ile Ala Leu Ser Leu Gly Leu Thr Pro Asn Phe Lys Ser Asn
530 535 540
Phe Asp Leu Ala Glu Asp Ala Lys Leu Gln Leu Ser Lys Asp Thr Tyr
545 550 555 560
Asp Asp Asp Leu Asp Asn Leu Leu Ala Gln Ile Gly Asp Gln Tyr Ala
565 570 575
Asp Leu Phe Leu Ala Ala Lys Asn Leu Ser Asp Ala Ile Leu Leu Ser
580 585 590
Asp Ile Leu Arg Val Asn Thr Glu Ile Thr Lys Ala Pro Leu Ser Ala
595 600 605
Ser Met Ile Lys Arg Tyr Asp Glu His His Gln Asp Leu Thr Leu Leu
610 615 620
Lys Ala Leu Val Arg Gln Gln Leu Pro Glu Lys Tyr Lys Glu Ile Phe
625 630 635 640
Phe Asp Gln Ser Lys Asn Gly Tyr Ala Gly Tyr Ile Asp Gly Gly Ala
645 650 655
Ser Gln Glu Glu Phe Tyr Lys Phe Ile Lys Pro Ile Leu Glu Lys Met
660 665 670
Asp Gly Thr Glu Glu Leu Leu Val Lys Leu Asn Arg Glu Asp Leu Leu
675 680 685
Arg Lys Gln Arg Thr Phe Asp Asn Gly Ser Ile Pro His Gln Ile His
690 695 700
Leu Gly Glu Leu His Ala Ile Leu Arg Arg Gln Glu Asp Phe Tyr Pro
705 710 715 720
Phe Leu Lys Asp Asn Arg Glu Lys Ile Glu Lys Ile Leu Thr Phe Arg
725 730 735
Ile Pro Tyr Tyr Val Gly Pro Leu Ala Arg Gly Asn Ser Arg Phe Ala
740 745 750
Trp Met Thr Arg Lys Ser Glu Glu Thr Ile Thr Pro Trp Asn Phe Glu
755 760 765
Glu Val Val Asp Lys Gly Ala Ser Ala Gln Ser Phe Ile Glu Arg Met
770 775 780
Thr Asn Phe Asp Lys Asn Leu Pro Asn Glu Lys Val Leu Pro Lys His
785 790 795 800
Ser Leu Leu Tyr Glu Tyr Phe Thr Val Tyr Asn Glu Leu Thr Lys Val
805 810 815
Lys Tyr Val Thr Glu Gly Met Arg Lys Pro Ala Phe Leu Ser Gly Glu
820 825 830
Gln Lys Lys Ala Ile Val Asp Leu Leu Phe Lys Thr Asn Arg Lys Val
835 840 845
Thr Val Lys Gln Leu Lys Glu Asp Tyr Phe Lys Lys Ile Glu Cys Phe
850 855 860
Asp Ser Val Glu Ile Ser Gly Val Glu Asp Arg Phe Asn Ala Ser Leu
865 870 875 880
Gly Thr Tyr His Asp Leu Leu Lys Ile Ile Lys Asp Lys Asp Phe Leu
885 890 895
Asp Asn Glu Glu Asn Glu Asp Ile Leu Glu Asp Ile Val Leu Thr Leu
900 905 910
Thr Leu Phe Glu Asp Arg Glu Met Ile Glu Glu Arg Leu Lys Thr Tyr
915 920 925
Ala His Leu Phe Asp Asp Lys Val Met Lys Gln Leu Lys Arg Arg Arg
930 935 940
Tyr Thr Gly Trp Gly Arg Leu Ser Arg Lys Leu Ile Asn Gly Ile Arg
945 950 955 960
Asp Lys Gln Ser Gly Lys Thr Ile Leu Asp Phe Leu Lys Ser Asp Gly
965 970 975
Phe Ala Asn Arg Asn Phe Met Gln Leu Ile His Asp Asp Ser Leu Thr
980 985 990
Phe Lys Glu Asp Ile Gln Lys Ala Gln Val Ser Gly Gln Gly Asp Ser
995 1000 1005
Leu His Glu His Ile Ala Asn Leu Ala Gly Ser Pro Ala Ile Lys
1010 1015 1020
Lys Gly Ile Leu Gln Thr Val Lys Val Val Asp Glu Leu Val Lys
1025 1030 1035
Val Met Gly Arg His Lys Pro Glu Asn Ile Val Ile Glu Met Ala
1040 1045 1050
Arg Glu Asn Gln Thr Thr Gln Lys Gly Gln Lys Asn Ser Arg Glu
1055 1060 1065
Arg Met Lys Arg Ile Glu Glu Gly Ile Lys Glu Leu Gly Ser Gln
1070 1075 1080
Ile Leu Lys Glu His Pro Val Glu Asn Thr Gln Leu Gln Asn Glu
1085 1090 1095
Lys Leu Tyr Leu Tyr Tyr Leu Gln Asn Gly Arg Asp Met Tyr Val
1100 1105 1110
Asp Gln Glu Leu Asp Ile Asn Arg Leu Ser Asp Tyr Asp Val Asp
1115 1120 1125
His Ile Val Pro Gln Ser Phe Leu Ala Asp Asp Ser Ile Asp Asn
1130 1135 1140
Lys Val Leu Thr Arg Ser Asp Lys Asn Arg Gly Lys Ser Asp Asn
1145 1150 1155
Val Pro Ser Glu Glu Val Val Lys Lys Met Lys Asn Tyr Trp Arg
1160 1165 1170
Gln Leu Leu Asn Ala Lys Leu Ile Thr Gln Arg Lys Phe Asp Asn
1175 1180 1185
Leu Thr Lys Ala Glu Arg Gly Gly Leu Ser Glu Leu Asp Lys Ala
1190 1195 1200
Gly Phe Ile Lys Arg Gln Leu Val Glu Thr Arg Gln Ile Thr Lys
1205 1210 1215
His Val Ala Gln Ile Leu Asp Ser Arg Met Asn Thr Lys Tyr Asp
1220 1225 1230
Glu Asn Asp Lys Leu Ile Arg Glu Val Lys Val Ile Thr Leu Lys
1235 1240 1245
Ser Lys Leu Val Ser Asp Phe Arg Lys Asp Phe Gln Phe Tyr Lys
1250 1255 1260
Val Arg Glu Ile Asn Asn Tyr His His Ala His Asp Ala Tyr Leu
1265 1270 1275
Asn Ala Val Val Gly Thr Ala Leu Ile Lys Lys Tyr Pro Ala Leu
1280 1285 1290
Glu Ser Glu Phe Val Tyr Gly Asp Tyr Lys Val Tyr Asp Val Arg
1295 1300 1305
Lys Met Ile Ala Lys Ser Glu Gln Glu Ile Gly Lys Ala Thr Ala
1310 1315 1320
Lys Tyr Phe Phe Tyr Ser Asn Ile Met Asn Phe Phe Lys Thr Glu
1325 1330 1335
Ile Thr Leu Ala Asn Gly Glu Ile Arg Lys Ala Pro Leu Ile Glu
1340 1345 1350
Thr Asn Gly Glu Thr Gly Glu Ile Val Trp Asp Lys Gly Arg Asp
1355 1360 1365
Phe Ala Thr Val Arg Lys Val Leu Ser Met Pro Gln Val Asn Ile
1370 1375 1380
Val Lys Lys Thr Glu Val Gln Thr Gly Gly Phe Ser Lys Glu Ser
1385 1390 1395
Ile Leu Pro Lys Arg Asn Ser Asp Lys Leu Ile Ala Arg Lys Lys
1400 1405 1410
Asp Trp Asp Pro Lys Lys Tyr Gly Gly Phe Asp Ser Pro Thr Val
1415 1420 1425
Ala Tyr Ser Val Leu Val Val Ala Lys Val Glu Lys Gly Lys Ser
1430 1435 1440
Lys Lys Leu Lys Ser Val Lys Glu Leu Leu Gly Ile Thr Ile Met
1445 1450 1455
Glu Arg Ser Ser Phe Glu Lys Asn Pro Ile Asp Phe Leu Glu Ala
1460 1465 1470
Lys Gly Tyr Lys Glu Val Lys Lys Asp Leu Ile Ile Lys Leu Pro
1475 1480 1485
Lys Tyr Ser Leu Phe Glu Leu Glu Asn Gly Arg Lys Arg Met Leu
1490 1495 1500
Ala Ser Ala Gly Glu Leu Gln Lys Gly Asn Glu Leu Ala Leu Pro
1505 1510 1515
Ser Lys Tyr Val Asn Phe Leu Tyr Leu Ala Ser His Tyr Glu Lys
1520 1525 1530
Leu Lys Gly Ser Pro Glu Asp Asn Glu Gln Lys Gln Leu Phe Val
1535 1540 1545
Glu Gln His Lys His Tyr Leu Asp Glu Ile Ile Glu Gln Ile Ser
1550 1555 1560
Glu Phe Ser Lys Arg Val Ile Leu Ala Asp Ala Asn Leu Asp Lys
1565 1570 1575
Val Leu Ser Ala Tyr Asn Lys His Arg Asp Lys Pro Ile Arg Glu
1580 1585 1590
Gln Ala Glu Asn Ile Ile His Leu Phe Thr Leu Thr Asn Leu Gly
1595 1600 1605
Ala Pro Ala Ala Phe Lys Tyr Phe Asp Thr Thr Ile Asp Arg Lys
1610 1615 1620
Arg Tyr Thr Ser Thr Lys Glu Val Leu Asp Ala Thr Leu Ile His
1625 1630 1635
Gln Ser Ile Thr Gly Leu Tyr Glu Thr Arg Ile Asp Leu Ser Gln
1640 1645 1650
Leu Gly Gly Asp Glu Phe Pro Lys Lys Lys Arg Lys Val Gly Gly
1655 1660 1665
Gly Gly Ser Pro Lys Lys Lys Arg Lys Val
1670 1675
<210> 51
<211> 20
<212> RNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
oligonucleotide"
<400> 51
gaguccgagc agaagaagaa 20
<210> 52
<211> 20
<212> RNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
oligonucleotide"
<400> 52
ggcacugcgg cuggaggugg 20
<210> 53
<211> 20
<212> RNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
oligonucleotide"
<400> 53
ggucacuuuu aacacaccca 20
<210> 54
<211> 55
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (34)..(39)
<223> a, t, g or c
<400> 54
tcgtcggcag cgtcagatgt gtataagaga cagnnnnnnc cccagtggct gctct 55
<210> 55
<211> 57
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (35)..(40)
<223> a, t, g or c
<400> 55
gtctcgtggg ctcggagatg tgtataagag acagnnnnnn ccaggcctcc ccaaagc 57
<210> 56
<211> 59
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (34)..(39)
<223> a, t, g or c
<400> 56
tcgtcggcag cgtcagatgt gtataagaga cagnnnnnng gaacccaggt agccagaga 59
<210> 57
<211> 57
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (35)..(40)
<223> a, t, g or c
<400> 57
gtctcgtggg ctcggagatg tgtataagag acagnnnnnn ggggtggggt cagacgt 57
<210> 58
<211> 59
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (34)..(39)
<223> a, t, g or c
<400> 58
tcgtcggcag cgtcagatgt gtataagaga cagnnnnnng cccattccct ctttagcca 59
<210> 59
<211> 61
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (35)..(40)
<223> a, t, g or c
<400> 59
gtctcgtggg ctcggagatg tgtataagag acagnnnnnn ggagcaggaa agtgaggtta 60
c 61
<210> 60
<211> 61
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (34)..(39)
<223> a, t, g or c
<400> 60
tcgtcggcag cgtcagatgt gtataagaga cagnnnnnna atggacacat gggtagtcag 60
g 61
<210> 61
<211> 64
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
primer"
<220>
<221> modified_base
<222> (35)..(40)
<223> a, t, g or c
<400> 61
gtctcgtggg ctcggagatg tgtataagag acagnnnnnn ggcttatatc caacacttcg 60
tggg 64
<210> 62
<211> 5076
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polynucleotide"
<400> 62
atggttagca aaggtgaaga actgtttaca ggtgttgttc cgattctggt tgaactggat 60
ggtgatgtta atggccacaa attttcagtt agcggtgaag gcgaaggtga tgcaacctat 120
ggtaaactga ccctgaaatt tatctgtacc accggcaaac tgccggttcc gtggccgaca 180
ctggttacca cactgaccta tggtgttcag tgttttagcc gttatccgga tcacatgaaa 240
cagcacgatt ttttcaaaag cgcaatgccg gaaggttatg ttcaagaacg taccatcttc 300
ttcaaagatg acggcaacta taaaacccgt gccgaagtta aatttgaagg tgataccctg 360
gtgaatcgca ttgaactgaa aggcatcgat tttaaagagg atggtaatat cctgggccac 420
aaactggaat ataattataa tagccacaac gtgtacatca tggccgacaa acagaaaaat 480
ggcatcaaag tgaacttcaa gatccgccat aatattgaag atggttcagt tcagctggcc 540
gatcattatc agcagaatac cccgattggt gatggtccgg ttctgctgcc ggataatcat 600
tatctgagca cccagagcaa actgagcaaa gatccgaatg aaaaacgtga tcacatggtg 660
ctgctggaat ttgttaccgc agcaggtatt accttaggta tggatgaact gtataaagtc 720
gacgcagaag cagcagcaaa agaagccgct gccaaagaag cggcagcgaa agaggcagcc 780
gcaaaagcac tggaagccga ggctgcggct aaagaggctg ctgcaaaaga agcagccgct 840
aaagaagctg cggctaaggc accggcagca aaacgtgtta aactggacgg tggtggtggt 900
agcaccggta tggacaagaa atacagcatc ggtttggata ttggcacgaa tagcgtgggt 960
tgggccgtta ttaccgacga gtacaaagtg ccgtccaaga aattcaaagt gctgggcaat 1020
accgatcgcc atagcatcaa gaaaaatctg attggcgcac tgctgttcga cagcggtgag 1080
actgccgaag ctacgcgtct gaagcgtacg gcgcgtcgtc gctacacccg ccgtaagaac 1140
cgtatttgct atctgcaaga aatcttcagc aacgaaatgg ccaaagttga tgatagcttt 1200
tttcaccgcc tggaagagag ctttctggtg gaagaggata agaaacacga gcgccatccg 1260
atttttggta acattgtcga tgaagtggca taccatgaga agtacccgac catctaccac 1320
cttcgtaaga aactggtgga cagcaccgat aaagctgatc tgcgtctgat ttacctggcg 1380
ctggcccaca tgattaagtt tcgcggtcat tttctgatcg agggcgatct gaatccggac 1440
aattctgatg ttgacaagct gtttattcaa cttgtacaga cctacaacca gttgttcgaa 1500
gagaacccga tcaatgcgag cggtgttgat gccaaagcaa ttctgagcgc acgcctgagc 1560
aaatctcgcc gtttggagaa cctgattgca cagctgccgg gtgagaagaa aaacggtctg 1620
ttcggcaatc tgattgcact gtccctgggc ttgaccccga attttaagag caacttcgac 1680
ctggccgaag atgcgaagct ccaattgagc aaagacacct acgacgatga cctggacaat 1740
ctgctggccc agattggcga ccagtacgca gatctgttct tggctgcgaa aaacctgagc 1800
gatgcaattc tgctgtcgga catcctgcgc gtgaatacgg aaatcacgaa agcgcctctg 1860
agcgcgtcta tgatcaagcg ctatgacgag caccaccaag atctgaccct gctgaaagct 1920
ctggtgagac aacaattgcc agagaagtat aaagaaattt tctttgacca gagcaaaaac 1980
ggctatgcgg gttacattga cggtggcgcc agccaagaag agttctacaa attcattaag 2040
cctatcctgg agaaaatgga tggcaccgaa gaactgctgg taaagctgaa tcgtgaagat 2100
ctgctgcgca aacagcgcac ttttgataac ggtagcattc cgcaccagat ccatctgggt 2160
gagttgcacg cgattttgcg tcgccaggaa gatttttatc cgttcttgaa agacaaccgt 2220
gagaaaatcg agaaaattct gacgttccgt atcccgtatt atgtcggccc gctggcgcgt 2280
ggtaatagcc gcttcgcgtg gatgacccgc aaatcagagg aaacgattac cccgtggaat 2340
tttgaggaag ttgttgataa gggtgcaagc gcgcagtcgt tcattgagcg tatgaccaac 2400
tttgacaaga atttgccgaa tgaaaaagtc ttgccgaagc actctctgct gtacgagtat 2460
tttaccgttt acaacgaatt gaccaaggtt aaatacgtca ccgaaggcat gcgcaaaccg 2520
gccttcctga gcggcgagca gaaaaaagca atcgttgacc tcttgtttaa gaccaaccgc 2580
aaggttacgg tcaaacaact gaaagaggac tatttcaaga aaattgaatg ttttgactcc 2640
gtagagatct ccggtgttga ggaccgtttc aacgcgagcc tgggcaccta ccatgatctg 2700
ctgaaaatta ttaaagacaa agattttctg gacaacgaag agaacgaaga tattctggaa 2760
gatatcgttc tgaccctgac gctgttcgaa gatcgtgaga tgattgagga acgtctgaaa 2820
acctacgcac acttgttcga tgacaaagtt atgaaacagc tgaagcgtcg tcgttacaca 2880
ggttggggcc gtctgagccg taagcttatc aatggtatcc gtgacaaaca gagcggtaag 2940
acgattctgg actttctgaa gtcagatggc ttcgccaatc gcaactttat gcaactgatt 3000
catgacgact ctctgacgtt caaggaagat atccaaaagg cacaggtgag cggtcagggt 3060
gatagcctgc atgagcatat cgcgaacctg gcgggtagcc cggctatcaa aaagggtatc 3120
ttacagactg tgaaagttgt ggatgaattg gttaaggtta tgggtcgtca caaaccggaa 3180
aatattgtga tcgagatggc acgtgaaaat cagacgacgc aaaagggtca aaaaaattct 3240
cgtgagcgca tgaaacgtat tgaagagggt atcaaagaat tgggcagcca aattctgaaa 3300
gaacacccgg tcgagaacac ccagctgcaa aacgaaaaac tgtatttata ctatctgcag 3360
aacggtcgtg acatgtacgt ggatcaagaa ctggacatca atcgtttgag cgattacgat 3420
gttgatcata ttgtgcctca gagctttctg aaagacgatt cgatcgacaa caaagtgctg 3480
acccgtagcg acaagaatcg tggtaagagc gataacgtgc cgagcgaaga agtcgttaag 3540
aaaatgaaaa actactggcg tcagctgctg aacgccaagc tgattaccca gcgtaagttc 3600
gataacctga cgaaagccga gcgtggaggc ctgagcgagc tggacaaggc cggctttatc 3660
aagcgtcaac tggtggaaac ccgtcagatc actaaacatg tggcacagat cctggactcc 3720
cgcatgaata cgaaatatga cgagaatgac aagttgatcc gtgaagtcaa agttattacg 3780
ctgaaaagca aactggtgtc cgatttccgt aaagacttcc agttctataa agtccgtgaa 3840
atcaacaact atcatcacgc ccacgatgcg tacttgaacg ctgttgtggg caccgcactg 3900
atcaagaaat accctaagct cgaaagcgag tttgtctatg gtgactataa agtttacgac 3960
gtgcgtaaga tgatcgccaa gagcgagcaa gaaattggta aggctaccgc aaagtacttt 4020
ttctacagca acatcatgaa cttcttcaaa accgagatta ccctggcgaa cggtgagatc 4080
cgtaaacggc cgctgattga gactaatggc gaaacgggcg agattgtgtg ggacaagggt 4140
cgcgatttcg ctacggttcg taaggtcctg agcatgccgc aagttaacat tgtcaagaaa 4200
actgaagtgc agacgggtgg ctttagcaaa gaatccatcc tgccgaagcg taatagcgat 4260
aaacttatcg cgcgtaaaaa agactgggac ccaaagaaat atggcggctt tgatagcccg 4320
accgtcgcgt atagcgtgtt agtggtcgcg aaagttgaaa agggcaagag caagaaactg 4380
aagtccgtca aagaacttct gggtatcacc atcatggaac gtagctcctt tgagaagaac 4440
ccgattgact tcttagaggc gaagggttat aaagaagtca aaaaagacct gattatcaag 4500
ctgccgaagt acagcctgtt tgagttggag aatggtcgta agcgcatgct ggcgagcgcg 4560
ggtgagctgc aaaagggcaa cgaactggcg ctgccgtcga aatacgtcaa ttttctgtac 4620
ctggccagcc actacgaaaa gctgaagggt tctccggaag ataacgaaca aaagcaactg 4680
ttcgttgagc aacataaaca ctacttggac gaaatcatcg agcaaattag cgaatttagc 4740
aaacgtgtca tcctggcgga cgcgaatctg gacaaggtcc tgtctgcata caataagcat 4800
cgcgacaaac caattcgtga gcaagcggag aatatcatcc acctgtttac gctgaccaac 4860
ctaggtgcgc cggcggcatt caagtatttc gatacgacca tcgaccgcaa gcgctatacc 4920
agcaccaaag aggtcctgga cgcgaccctg atccaccaga gcattaccgg cttatacgaa 4980
acccgtattg atttgagcca actgggtggc gatgaattcc cgaaaaaaaa gcgcaaagtt 5040
ggtggcggtg gtagcccgaa aaagaaacgt aaagtg 5076
<210> 63
<211> 5013
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polynucleotide"
<400> 63
atggttagca aaggtgaagc cgtgattaaa gaatttatgc gctttaaggt tcacatggaa 60
ggtagcatga atggccatga atttgaaatt gaaggtgaag gcgaaggtcg tccgtatgaa 120
ggcacccaga ccgcaaaact gaaagttacc aaaggtggtc cgctgccgtt tagctgggat 180
attctgagtc cgcagtttat gtatggtagc cgtgcattta ccaaacatcc ggcagatatt 240
ccggattatt acaaacagag ctttccggaa ggttttaaat gggaacgtgt gatgaatttt 300
gaagatggtg gtgcagttac cgttacacag gataccagcc tggaagatgg caccctgatc 360
tataaagtta aactgcgtgg caccaatttt ccgcctgatg gtccggttat gcagaaaaaa 420
acaatgggtt gggaagcaag caccgaacgt ctgtatcctg aagatggcgt tctgaaaggt 480
gatatcaaaa tggcactgcg tctgaaagat ggcggtcgtt atctggcaga tttcaaaacc 540
acctataaag ccaaaaaacc tgttcagatg cctggtgcct ataatgttga tcgtaaactg 600
gatattacca gccacaacga agattatacc gttgtggaac agtatgaacg tagcgaaggc 660
cgtcatagca caggtggtat ggatgaactg tataaagtcg acagcggtgg tagcagcggt 720
ggttcaagcg gtagcgaaac accgggtaca agcgaaagcg caacaccgga aagcagtggt 780
ggtagttcag gtggtagtcc ggcagcaaaa cgtgtgaaac tggatggcgg tggcggtagc 840
accggtatgg acaagaaata cagcatcggt ttggatattg gcacgaatag cgtgggttgg 900
gccgttatta ccgacgagta caaagtgccg tccaagaaat tcaaagtgct gggcaatacc 960
gatcgccata gcatcaagaa aaatctgatt ggcgcactgc tgttcgacag cggtgagact 1020
gccgaagcta cgcgtctgaa gcgtacggcg cgtcgtcgct acacccgccg taagaaccgt 1080
atttgctatc tgcaagaaat cttcagcaac gaaatggcca aagttgatga tagctttttt 1140
caccgcctgg aagagagctt tctggtggaa gaggataaga aacacgagcg ccatccgatt 1200
tttggtaaca ttgtcgatga agtggcatac catgagaagt acccgaccat ctaccacctt 1260
cgtaagaaac tggtggacag caccgataaa gctgatctgc gtctgattta cctggcgctg 1320
gcccacatga ttaagtttcg cggtcatttt ctgatcgagg gcgatctgaa tccggacaat 1380
tctgatgttg acaagctgtt tattcaactt gtacagacct acaaccagtt gttcgaagag 1440
aacccgatca atgcgagcgg tgttgatgcc aaagcaattc tgagcgcacg cctgagcaaa 1500
tctcgccgtt tggagaacct gattgcacag ctgccgggtg agaagaaaaa cggtctgttc 1560
ggcaatctga ttgcactgtc cctgggcttg accccgaatt ttaagagcaa cttcgacctg 1620
gccgaagatg cgaagctcca attgagcaaa gacacctacg acgatgacct ggacaatctg 1680
ctggcccaga ttggcgacca gtacgcagat ctgttcttgg ctgcgaaaaa cctgagcgat 1740
gcaattctgc tgtcggacat cctgcgcgtg aatacggaaa tcacgaaagc gcctctgagc 1800
gcgtctatga tcaagcgcta tgacgagcac caccaagatc tgaccctgct gaaagctctg 1860
gtgagacaac aattgccaga gaagtataaa gaaattttct ttgaccagag caaaaacggc 1920
tatgcgggtt acattgacgg tggcgccagc caagaagagt tctacaaatt cattaagcct 1980
atcctggaga aaatggatgg caccgaagaa ctgctggtaa agctgaatcg tgaagatctg 2040
ctgcgcaaac agcgcacttt tgataacggt agcattccgc accagatcca tctgggtgag 2100
ttgcacgcga ttttgcgtcg ccaggaagat ttttatccgt tcttgaaaga caaccgtgag 2160
aaaatcgaga aaattctgac gttccgtatc ccgtattatg tcggcccgct ggcgcgtggt 2220
aatagccgct tcgcgtggat gacccgcaaa tcagaggaaa cgattacccc gtggaatttt 2280
gaggaagttg ttgataaggg tgcaagcgcg cagtcgttca ttgagcgtat gaccaacttt 2340
gacaagaatt tgccgaatga aaaagtcttg ccgaagcact ctctgctgta cgagtatttt 2400
accgtttaca acgaattgac caaggttaaa tacgtcaccg aaggcatgcg caaaccggcc 2460
ttcctgagcg gcgagcagaa aaaagcaatc gttgacctct tgtttaagac caaccgcaag 2520
gttacggtca aacaactgaa agaggactat ttcaagaaaa ttgaatgttt tgactccgta 2580
gagatctccg gtgttgagga ccgtttcaac gcgagcctgg gcacctacca tgatctgctg 2640
aaaattatta aagacaaaga ttttctggac aacgaagaga acgaagatat tctggaagat 2700
atcgttctga ccctgacgct gttcgaagat cgtgagatga ttgaggaacg tctgaaaacc 2760
tacgcacact tgttcgatga caaagttatg aaacagctga agcgtcgtcg ttacacaggt 2820
tggggccgtc tgagccgtaa gcttatcaat ggtatccgtg acaaacagag cggtaagacg 2880
attctggact ttctgaagtc agatggcttc gccaatcgca actttatgca actgattcat 2940
gacgactctc tgacgttcaa ggaagatatc caaaaggcac aggtgagcgg tcagggtgat 3000
agcctgcatg agcatatcgc gaacctggcg ggtagcccgg ctatcaaaaa gggtatctta 3060
cagactgtga aagttgtgga tgaattggtt aaggttatgg gtcgtcacaa accggaaaat 3120
attgtgatcg agatggcacg tgaaaatcag acgacgcaaa agggtcaaaa aaattctcgt 3180
gagcgcatga aacgtattga agagggtatc aaagaattgg gcagccaaat tctgaaagaa 3240
cacccggtcg agaacaccca gctgcaaaac gaaaaactgt atttatacta tctgcagaac 3300
ggtcgtgaca tgtacgtgga tcaagaactg gacatcaatc gtttgagcga ttacgatgtt 3360
gatcatattg tgcctcagag ctttctgaaa gacgattcga tcgacaacaa agtgctgacc 3420
cgtagcgaca agaatcgtgg taagagcgat aacgtgccga gcgaagaagt cgttaagaaa 3480
atgaaaaact actggcgtca gctgctgaac gccaagctga ttacccagcg taagttcgat 3540
aacctgacga aagccgagcg tggaggcctg agcgagctgg acaaggccgg ctttatcaag 3600
cgtcaactgg tggaaacccg tcagatcact aaacatgtgg cacagatcct ggactcccgc 3660
atgaatacga aatatgacga gaatgacaag ttgatccgtg aagtcaaagt tattacgctg 3720
aaaagcaaac tggtgtccga tttccgtaaa gacttccagt tctataaagt ccgtgaaatc 3780
aacaactatc atcacgccca cgatgcgtac ttgaacgctg ttgtgggcac cgcactgatc 3840
aagaaatacc ctaagctcga aagcgagttt gtctatggtg actataaagt ttacgacgtg 3900
cgtaagatga tcgccaagag cgagcaagaa attggtaagg ctaccgcaaa gtactttttc 3960
tacagcaaca tcatgaactt cttcaaaacc gagattaccc tggcgaacgg tgagatccgt 4020
aaacggccgc tgattgagac taatggcgaa acgggcgaga ttgtgtggga caagggtcgc 4080
gatttcgcta cggttcgtaa ggtcctgagc atgccgcaag ttaacattgt caagaaaact 4140
gaagtgcaga cgggtggctt tagcaaagaa tccatcctgc cgaagcgtaa tagcgataaa 4200
cttatcgcgc gtaaaaaaga ctgggaccca aagaaatatg gcggctttga tagcccgacc 4260
gtcgcgtata gcgtgttagt ggtcgcgaaa gttgaaaagg gcaagagcaa gaaactgaag 4320
tccgtcaaag aacttctggg tatcaccatc atggaacgta gctcctttga gaagaacccg 4380
attgacttct tagaggcgaa gggttataaa gaagtcaaaa aagacctgat tatcaagctg 4440
ccgaagtaca gcctgtttga gttggagaat ggtcgtaagc gcatgctggc gagcgcgggt 4500
gagctgcaaa agggcaacga actggcgctg ccgtcgaaat acgtcaattt tctgtacctg 4560
gccagccact acgaaaagct gaagggttct ccggaagata acgaacaaaa gcaactgttc 4620
gttgagcaac ataaacacta cttggacgaa atcatcgagc aaattagcga atttagcaaa 4680
cgtgtcatcc tggcggacgc gaatctggac aaggtcctgt ctgcatacaa taagcatcgc 4740
gacaaaccaa ttcgtgagca agcggagaat atcatccacc tgtttacgct gaccaaccta 4800
ggtgcgccgg cggcattcaa gtatttcgat acgaccatcg accgcaagcg ctataccagc 4860
accaaagagg tcctggacgc gaccctgatc caccagagca ttaccggctt atacgaaacc 4920
cgtattgatt tgagccaact gggtggcgat gaattcccga aaaaaaagcg caaagttggt 4980
ggcggtggta gcccgaaaaa gaaacgtaaa gtg 5013
<210> 64
<211> 5034
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polynucleotide"
<400> 64
atggttagca aaggtgaaga actgtttaca ggtgttgttc cgattctggt tgaactggat 60
ggtgatgtta atggccacaa attttcagtt agcggtgaag gcgaaggtga tgcaacctat 120
ggtaaactga ccctgaaatt tatctgtacc accggcaaac tgccggttcc gtggccgaca 180
ctggttacca cactgaccta tggtgttcag tgttttagcc gttatccgga tcacatgaaa 240
cagcacgatt ttttcaaaag cgcaatgccg gaaggttatg ttcaagaacg taccatcttc 300
ttcaaagatg acggcaacta taaaacccgt gccgaagtta aatttgaagg tgataccctg 360
gtgaatcgca ttgaactgaa aggcatcgat tttaaagagg atggtaatat cctgggccac 420
aaactggaat ataattataa tagccacaac gtgtacatca tggccgacaa acagaaaaat 480
ggcatcaaag tgaacttcaa gatccgccat aatattgaag atggttcagt tcagctggcc 540
gatcattatc agcagaatac cccgattggt gatggtccgg ttctgctgcc ggataatcat 600
tatctgagca cccagagcaa actgagcaaa gatccgaatg aaaaacgtga tcacatggtg 660
ctgctggaat ttgttaccgc agcaggtatt accttaggta tggatgaact gtataaagtc 720
gacagcggtg gtagcagcgg tggttcaagc ggtagcgaaa caccgggtac aagcgaaagc 780
gcaacaccgg aaagcagtgg tggtagctca ggtggtagtc cggcagcaaa acgtgttaaa 840
ctggacggtg gtggtggtag caccggtatg gacaagaaat acagcatcgg tttggatatt 900
ggcacgaata gcgtgggttg ggccgttatt accgacgagt acaaagtgcc gtccaagaaa 960
ttcaaagtgc tgggcaatac cgatcgccat agcatcaaga aaaatctgat tggcgcactg 1020
ctgttcgaca gcggtgagac tgccgaagct acgcgtctga agcgtacggc gcgtcgtcgc 1080
tacacccgcc gtaagaaccg tatttgctat ctgcaagaaa tcttcagcaa cgaaatggcc 1140
aaagttgatg atagcttttt tcaccgcctg gaagagagct ttctggtgga agaggataag 1200
aaacacgagc gccatccgat ttttggtaac attgtcgatg aagtggcata ccatgagaag 1260
tacccgacca tctaccacct tcgtaagaaa ctggtggaca gcaccgataa agctgatctg 1320
cgtctgattt acctggcgct ggcccacatg attaagtttc gcggtcattt tctgatcgag 1380
ggcgatctga atccggacaa ttctgatgtt gacaagctgt ttattcaact tgtacagacc 1440
tacaaccagt tgttcgaaga gaacccgatc aatgcgagcg gtgttgatgc caaagcaatt 1500
ctgagcgcac gcctgagcaa atctcgccgt ttggagaacc tgattgcaca gctgccgggt 1560
gagaagaaaa acggtctgtt cggcaatctg attgcactgt ccctgggctt gaccccgaat 1620
tttaagagca acttcgacct ggccgaagat gcgaagctcc aattgagcaa agacacctac 1680
gacgatgacc tggacaatct gctggcccag attggcgacc agtacgcaga tctgttcttg 1740
gctgcgaaaa acctgagcga tgcaattctg ctgtcggaca tcctgcgcgt gaatacggaa 1800
atcacgaaag cgcctctgag cgcgtctatg atcaagcgct atgacgagca ccaccaagat 1860
ctgaccctgc tgaaagctct ggtgagacaa caattgccag agaagtataa agaaattttc 1920
tttgaccaga gcaaaaacgg ctatgcgggt tacattgacg gtggcgccag ccaagaagag 1980
ttctacaaat tcattaagcc tatcctggag aaaatggatg gcaccgaaga actgctggta 2040
aagctgaatc gtgaagatct gctgcgcaaa cagcgcactt ttgataacgg tagcattccg 2100
caccagatcc atctgggtga gttgcacgcg attttgcgtc gccaggaaga tttttatccg 2160
ttcttgaaag acaaccgtga gaaaatcgag aaaattctga cgttccgtat cccgtattat 2220
gtcggcccgc tggcgcgtgg taatagccgc ttcgcgtgga tgacccgcaa atcagaggaa 2280
acgattaccc cgtggaattt tgaggaagtt gttgataagg gtgcaagcgc gcagtcgttc 2340
attgagcgta tgaccaactt tgacaagaat ttgccgaatg aaaaagtctt gccgaagcac 2400
tctctgctgt acgagtattt taccgtttac aacgaattga ccaaggttaa atacgtcacc 2460
gaaggcatgc gcaaaccggc cttcctgagc ggcgagcaga aaaaagcaat cgttgacctc 2520
ttgtttaaga ccaaccgcaa ggttacggtc aaacaactga aagaggacta tttcaagaaa 2580
attgaatgtt ttgactccgt agagatctcc ggtgttgagg accgtttcaa cgcgagcctg 2640
ggcacctacc atgatctgct gaaaattatt aaagacaaag attttctgga caacgaagag 2700
aacgaagata ttctggaaga tatcgttctg accctgacgc tgttcgaaga tcgtgagatg 2760
attgaggaac gtctgaaaac ctacgcacac ttgttcgatg acaaagttat gaaacagctg 2820
aagcgtcgtc gttacacagg ttggggccgt ctgagccgta agcttatcaa tggtatccgt 2880
gacaaacaga gcggtaagac gattctggac tttctgaagt cagatggctt cgccaatcgc 2940
aactttatgc aactgattca tgacgactct ctgacgttca aggaagatat ccaaaaggca 3000
caggtgagcg gtcagggtga tagcctgcat gagcatatcg cgaacctggc gggtagcccg 3060
gctatcaaaa agggtatctt acagactgtg aaagttgtgg atgaattggt taaggttatg 3120
ggtcgtcaca aaccggaaaa tattgtgatc gagatggcac gtgaaaatca gacgacgcaa 3180
aagggtcaaa aaaattctcg tgagcgcatg aaacgtattg aagagggtat caaagaattg 3240
ggcagccaaa ttctgaaaga acacccggtc gagaacaccc agctgcaaaa cgaaaaactg 3300
tatttatact atctgcagaa cggtcgtgac atgtacgtgg atcaagaact ggacatcaat 3360
cgtttgagcg attacgatgt tgatcatatt gtgcctcaga gctttctggc ggacgattcg 3420
atcgacaaca aagtgctgac ccgtagcgac aagaatcgtg gtaagagcga taacgtgccg 3480
agcgaagaag tcgttaagaa aatgaaaaac tactggcgtc agctgctgaa cgccaagctg 3540
attacccagc gtaagttcga taacctgacg aaagccgagc gtggaggcct gagcgagctg 3600
gacaaggccg gctttatcaa gcgtcaactg gtggaaaccc gtcagatcac taaacatgtg 3660
gcacagatcc tggactcccg catgaatacg aaatatgacg agaatgacaa gttgatccgt 3720
gaagtcaaag ttattacgct gaaaagcaaa ctggtgtccg atttccgtaa agacttccag 3780
ttctataaag tccgtgaaat caacaactat catcacgccc acgatgcgta cttgaacgct 3840
gttgtgggca ccgcactgat caagaaatac cctgcactcg aaagcgagtt tgtctatggt 3900
gactataaag tttacgacgt gcgtaagatg atcgccaaga gcgagcaaga aattggtaag 3960
gctaccgcaa agtacttttt ctacagcaac atcatgaact tcttcaaaac cgagattacc 4020
ctggcgaacg gtgagatccg taaagcgccg ctgattgaga ctaatggcga aacgggcgag 4080
attgtgtggg acaagggtcg cgatttcgct acggttcgta aggtcctgag catgccgcaa 4140
gttaacattg tcaagaaaac tgaagtgcag acgggtggct ttagcaaaga atccatcctg 4200
ccgaagcgta atagcgataa acttatcgcg cgtaaaaaag actgggaccc aaagaaatat 4260
ggcggctttg atagcccgac cgtcgcgtat agcgtgttag tggtcgcgaa agttgaaaag 4320
ggcaagagca agaaactgaa gtccgtcaaa gaacttctgg gtatcaccat catggaacgt 4380
agctcctttg agaagaaccc gattgacttc ttagaggcga agggttataa agaagtcaaa 4440
aaagacctga ttatcaagct gccgaagtac agcctgtttg agttggagaa tggtcgtaag 4500
cgcatgctgg cgagcgcggg tgagctgcaa aagggcaacg aactggcgct gccgtcgaaa 4560
tacgtcaatt ttctgtacct ggccagccac tacgaaaagc tgaagggttc tccggaagat 4620
aacgaacaaa agcaactgtt cgttgagcaa cataaacact acttggacga aatcatcgag 4680
caaattagcg aatttagcaa acgtgtcatc ctggcggacg cgaatctgga caaggtcctg 4740
tctgcataca ataagcatcg cgacaaacca attcgtgagc aagcggagaa tatcatccac 4800
ctgtttacgc tgaccaacct aggtgcgccg gcggcattca agtatttcga tacgaccatc 4860
gaccgcaagc gctataccag caccaaagag gtcctggacg cgaccctgat ccaccagagc 4920
attaccggct tatacgaaac ccgtattgat ttgagccaac tgggtggcga tgaattcccg 4980
aaaaaaaagc gcaaagttgg tggcggtggt agcccgaaaa agaaacgtaa agtg 5034
<210> 65
<211> 20
<212> RNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
oligonucleotide"
<400> 65
gcucccauca caucaaccgg 20
<210> 66
<211> 20
<212> RNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
oligonucleotide"
<400> 66
ggugagugag ugugugcgug 20
Claims (37)
- 적어도 하나의 마커 단백질에 연결된 Cas9 단백질을 포함하는 융합 단백질.
- 청구항 1에 있어서, 적어도 하나의 마커 단백질은 화학 결합에 의해 직접적으로, 링커에 의해 간접적으로, 또는 이의 조합으로 Cas9 단백질에 연결되는, 융합 단백질.
- 청구항 1에 있어서, 적어도 하나의 마커 단백질은 직접적으로 또는 링커에 의해 간접적으로 Cas9 단백질에 연결되는, 융합 단백질.
- 청구항 1 내지 3 중 어느 한 항에 있어서, 적어도 하나의 마커 단백질은 N-말단, C-말단, 내부 위치, 또는 이의 조합에서 Cas9 단백질에 연결되는, 융합 단백질.
- 청구항 1 내지 4 중 어느 한 항에 있어서, 적어도 하나의 핵 위치화 신호를 추가로 포함하는, 융합 단백질.
- 청구항 5에 있어서, 핵 위치화 신호, 마커 단백질, 링커 및 Cas9 단백질은 (N-말단에서 C-말단으로) 다음 순서로 배열되는, 융합 단백질:
마커 단백질 - 링커 - 핵 위치화 신호 - Cas9 단백질;
마커 단백질 - 핵 위치화 신호 - 링커 - Cas9 단백질;
핵 위치화 신호 - 링커 - 마커 단백질 - Cas9 단백질;
핵 위치화 신호 - 마커 단백질 - 링커 - Cas9 단백질;
마커 단백질 - 링커 - 핵 위치화 신호 - 링커 - Cas9 단백질; 또는
핵 위치화 신호 - 링커 - 마커 단백질 - 링커 - Cas9 단백질. - 청구항 6에 있어서, 핵 위치화 신호, 마커 단백질, 링커 및 Cas9 단백질은 (N-말단에서 C-말단으로) 다음 순서로 배열되는, 융합 단백질:
마커 단백질 - 링커 - 핵 위치화 신호 - Cas9 단백질. - 청구항 1 내지 7 중 어느 한 항에 있어서, 적어도 하나의 이종 도메인을 추가로 포함하는, 융합 단백질.
- 청구항 8에 있어서, 적어도 하나의 이종 도메인은 세포-침투 도메인, 염색질 조절 모티프, 후성 변형 도메인, 전사 조절 도메인, RNA 압타머 결합 도메인, 또는 이의 조합인, 융합 단백질.
- 청구항 1 내지 9중 어느 한 항에 있어서, 융합 단백질은, 뉴클레아제이고 이중 가닥 서열의 두 가닥 모두를 절단하거나, 니카아제이고 이중 가닥 서열 중 한 가닥을 절단하거나, 뉴클레아제 또는 니카아제 활성을 가지지 않는, 융합 단백질.
- 청구항 1 내지 10 중 어느 한 항에 있어서, cas9 단백질은 스트렙토코쿠스 피오게네스 (Streptococcus pyogenes), 스트렙토코쿠스 써모필루스 (Streptococcus thermophilus), 네이세리아 메닝기티디스 (Neisseria meningitidis), 스타필로코쿠스 아우레우스 (Staphylococcus aureus), 또는 캄필로박터 제주니 (Campylobacter jejuni)에서 유래하는, 융합 단백질.
- 청구항 1 내지 11 중 어느 한 항에 있어서, 마커 단백질은 서열 번호: 19 또는 20과 적어도 90% 서열 동일성을 가지는 아미노산 서열을 가지는, 융합 단백질.
- 청구항 1 내지 11 중 어느 한 항에 있어서, 마커 단백질은 서열 번호: 19 또는 20에 제시된 아미노산 서열을 가지는, 융합 단백질.
- 청구항 3 내지 13 중 어느 한 항에 있어서, 링커는 서열 번호: 35 또는 36과 적어도 90% 서열 동일성을 가지는 아미노산 서열을 가지는, 융합 단백질.
- 청구항 3 내지 13 중 어느 한 항에 있어서, 링커는 서열 번호: 35 또는 36에 제시된 아미노산 서열을 가지는, 융합 단백질.
- 청구항 3 내지 15 중 어느 한 항에 있어서, 융합 단백질은 서열 번호: 48, 49, 또는 50과 적어도 90% 서열 동일성을 가지는 아미노산 서열을 가지는, 융합 단백질.
- 청구항 3 내지 15 중 어느 한 항에 있어서, 융합 단백질은 서열 번호: 48, 49, 또는 50에 제시된 아미노산 서열을 가지는, 융합 단백질.
- 청구항 1 내지 17 중 어느 한 항의 융합 단백질 및 조작된 가이드 RNA를 포함하는 시스템.
- 청구항 18에 있어서, 조작된 가이드 RNA는 단일 분자인, 시스템.
- 청구항 18 내지 19 중 어느 한 항에 있어서, 조작된 가이드 RNA 서열은 진핵 세포내의 조작된 가이드 RNA 내부에서 염기 페어링을 촉진하거나, 조작된 가이드 RNA 내부에서 염기 페어링을 최소화하거나, 조작된 가이드 RNA의 안정성을 증가시키거나, 조작된 가이드 RNA의 전사를 촉진하거나, 또는 이의 조합인, 시스템.
- 청구항 1 내지 17 중 어느 한 항의 융합 단백질을 인코딩하는 복수의 핵산.
- 청구항 18 내지 20 중 어느 한 항의 시스템을 인코딩하는 복수의 핵산으로서, 이러한 복수의 핵산은 융합 단백질을 인코딩하는 적어도 하나의 핵산, 및 조작된 가이드 RNA를 인코딩하는 적어도 하나의 핵산을 포함하는, 복수의 핵산.
- 청구항 14에 있어서, 융합 단백질을 인코딩하는 적어도 하나의 핵산은 RNA인, 복수의 핵산.
- 청구항 14에 있어서, 융합 단백질을 인코딩하는 적어도 하나의 핵산은 DNA인, 복수의 핵산.
- 청구항 21 내지 24 중 어느 한 항에 있어서, 융합 단백질을 인코딩하는 적어도 하나의 핵산은 진핵 세포에서의 발현에 코돈 최적화되는, 복수의 핵산.
- 청구항 25에 있어서, 진핵 세포는 인간 세포, 비-인간 포유동물 세포, 비-포유동물 척추동물 세포, 무척추동물 세포, 식물 세포, 또는 단세포 진핵 유기체인, 복수의 핵산.
- 청구항 22 내지 26 중 어느 한 항에 있어서, 조작된 가이드 RNA를 인코딩하는 적어도 하나의 핵산은 DNA인, 복수의 핵산.
- 청구항 22내지 27 중 어느 한 항에 있어서, 융합 단백질을 인코딩하는 적어도 하나의 핵산은 시험관내 RNA 합성 또는 박테리아 세포에서 단백질 발현을 위해 파지 프로모터 서열에 작동가능하게 연결되고, 상기 조작된 가이드 RNA를 인코딩하는 적어도 하나의 핵산은 시험관내 RNA 합성을 위해 파지 프로모터 서열에 작동가능하게 연결되는, 복수의 핵산.
- 청구항 22 내지 27 중 어느 한 항에 있어서, 융합 단백질을 인코딩하는 적어도 하나의 핵산은 진핵 세포에서 발현을 위해 진핵 프로모터 서열에 작동가능하게 연결되고, 상기 조작된 가이드 RNA를 인코딩하는 적어도 하나의 핵산은 진핵 세포에서 발현을 위해 진핵 프로모터 서열에 작동가능하게 연결되는, 복수의 핵산.
- 청구항 21 내지 29 중 어느 한 항의 복수의 핵산을 포함하는 적어도 하나의 벡터.
- 청구항 30에 있어서, 플라스미드 벡터, 바이러스 벡터, 또는 자가-복제 바이러스 RNA 레플리콘인, 적어도 하나의 벡터.
- 청구항 1 내지 17에 정의된 융합 단백질, 청구항 18 내지 20에 정의된 시스템, 청구항 21 내지 29에 정의된 복수의 핵산, 또는 청구항 30 내지 31에 정의된 적어도 하나의 벡터를 포함하는 적어도 하나의 시스템을 포함하는 진핵 세포.
- 청구항 32에 있어서, 인간 세포, 비-인간 포유동물 세포, 식물 세포, 비-포유동물 척추동물 세포, 무척추동물 세포, 또는 단세포 진핵 유기체인, 진핵 세포.
- 청구항 32 내지 33 중 어느 한 항에 있어서, 생체내, 생체외, 또는 시험관내인, 진핵 세포.
- 청구항 1 내지 31 중 어느 한 항의 융합 단백질, 시스템, 복수의 핵산, 또는 벡터를 살아있는 또는 화학적으로 고정된 진핵 세포에 도입하는 단계 및 마커 단백질로부터의 신호를 탐지하는 단계를 포함하는, 살아있는 진핵 세포 또는 화학적으로 고정된 진핵 세포 내에서 염색체 동일성 및 위치를 결정하는 방법.
- 청구항 35에 있어서, 진핵 세포는 인간 세포, 비-인간 포유동물 세포, 식물 세포, 비-포유동물 척추동물 세포, 무척추동물 세포, 또는 단세포 진핵 유기체인, 방법.
- 청구항 35 내지 36 중 어느 한 항에 있어서, 진핵 세포는 생체내, 생체외 또는 시험관내에 존재하는, 방법.
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| WO2023039424A2 (en) * | 2021-09-08 | 2023-03-16 | Flagship Pioneering Innovations Vi, Llc | Methods and compositions for modulating a genome |
| US20230265405A1 (en) * | 2022-02-22 | 2023-08-24 | Massachusetts Institute Of Technology | Engineered nucleases and methods of use thereof |
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2020
- 2020-02-13 CA CA3129835A patent/CA3129835A1/en active Pending
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- 2020-02-13 AU AU2020221274A patent/AU2020221274B2/en active Active
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- 2020-02-13 BR BR112021016019-1A patent/BR112021016019A2/pt unknown
- 2020-02-13 US US16/790,399 patent/US10947517B2/en active Active
- 2020-02-13 KR KR1020217029732A patent/KR20210139271A/ko not_active Application Discontinuation
- 2020-02-13 CN CN202080029346.8A patent/CN113728099A/zh active Pending
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2024
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| Publication number | Publication date |
|---|---|
| JP2023156365A (ja) | 2023-10-24 |
| US20210163910A1 (en) | 2021-06-03 |
| US10947517B2 (en) | 2021-03-16 |
| IL285300A (en) | 2021-09-30 |
| EP3924475A1 (en) | 2021-12-22 |
| US11965184B2 (en) | 2024-04-23 |
| WO2020168102A1 (en) | 2020-08-20 |
| CN113728099A (zh) | 2021-11-30 |
| CA3129835A1 (en) | 2020-08-20 |
| SG11202108812SA (en) | 2021-09-29 |
| BR112021016019A2 (pt) | 2021-10-05 |
| US20230287377A1 (en) | 2023-09-14 |
| AU2020221274A1 (en) | 2021-08-26 |
| JP2022520104A (ja) | 2022-03-28 |
| US20200263155A1 (en) | 2020-08-20 |
| AU2020221274B2 (en) | 2024-02-08 |
| AU2024202275A1 (en) | 2024-05-23 |
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