KR20230159550A - 유도 만능 줄기 세포 수득 방법 - Google Patents
유도 만능 줄기 세포 수득 방법 Download PDFInfo
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Abstract
본원에서는 조혈 계통의 세포로부터 유도 만능 줄기 세포를 수득하는 방법을 제공한다.
Description
서열 목록
본 출원은 ASCII 포맷으로 전자적으로 제출된 서열 목록을 포함하고, 그의 전문이 본원에서 참조로 포함된다. 2022년 3월 23일 작성된 상기 ASCII 사본의 명칭은 025450_WO014_SL.txt이고, 그 크기는 62,969 바이트이다.
세포 요법은 다양한 질환 및 병태의 치료에 대한 큰 가능성을 제공한다. 세포 요법에서, 자가 또는 동종 세포를 환자에게 이식하여 유전 장애, 암, 신경 장애, 심장 장애 또는 눈 관련 문제를 비롯한 다양한 의학적 상태로 인해 발생할 수 있는 결함이 있거나, 손상된 조직 또는 세포를 교체하거나, 또는 수복시킨다. 체세포에서 생성된 만능 줄기 세포를 비롯한 만능 줄기 세포는 세포 요법에 특히 유용하다. K. 타카하시(K. Takahashi) 및 S. 야마나카(S. Yamanaka)의 중대한 작업을 통해 4개의 재프로그래밍 전사 인자, Oct3/4, Klf4, Sox2, 및 c-Myc를 발현하는 레트로바이러스 벡터가 형질도입된 마우스 섬유아세포로부터의 만능 줄기 세포의 유도가 입증되었다 (Cell (2006) 126:663-76). 그러나, 레트로바이러스 벡터는 숙주 게놈에 삽입 돌연변이를 유발할 수 있으며, 이에 임상 환경에서 사용하기에 이상적인 벡터는 아니다.
따라서, 재프로그래밍 인자를 체세포에 도입하기 위해 RNA 기반 접근법이 시도되어 왔다. 그러한 접근법 중 하나는 알파바이러스 기반 바이러스 RNA 레플리콘을 이용한다. 토가비리다에(Togaviridae) 과의 30개 초과의 바이러스 속을 구성하는 알파바이러스는 지질 외피 보유 (+) 센스 RNA 바이러스이다. 뉴 월드 알파바이러스로는 동부, 서부 및 베네수엘라 말 뇌염 바이러스 (각각 EEEV, WEEV 및 VEEV)를 포함하고, 북미 및 남미에서 발견된다. 올드 월드 알파바이러스로는 치쿤군야(chikungunya: CHIK), 신드비스(Sindbis), 로스 리버(Ross River) 및 오뇽뇽(O'nyong-nyong) 바이러스를 포함한다.
알파바이러스는 대략 14 kb 길이의 (+) 센스 단일 가닥 RNA 게놈을 함유한다. 숙주 세포 내로의 진입 후, 알파바이러스 입자는 분해되어 게놈 RNA를 세포의 세포질로 방출한다. 바이러스 게놈의 번역을 통해 비구조 폴리단백질인 P1234가 생성되고, 이어서, 이는 프로테아제에 의해 절단되어 비구조 단백질 (nsP1, nsP2, nsP3, 및 nsP4)을 생성한다. 비구조 단백질은 RNA 복제 및 전사에 관여한다. 서브게놈 RNA - 26S RNA - 또한 전사를 통해 바이러스 게놈으로부터 생성된다. 26S RNA 번역으로 구조 폴리단백질이 생성되고, 이는 절단되어 구조 단백질 (예컨대, VEEV의 경우, C, E3, E2, 6K, 및 E1)을 생성한다. 구조 단백질은 발아 및 바이러스 캡시드화에 관여한다. 예컨대, 문헌 [Shin et al., PNAS (2012) 109(41):16534-9]; [Jose et al., Future Microbiol. (2009) 4:837-56; [Hardy and Strauss, J Virol . (1989) 63(11):4653-64; [Melancon and Garoff, J Virol . (1987) 61(5):1301-9; 및 [Strauss et al., Virology (1984) 133(1):92-110]; 및 [Glanville et al., PNAS (1976) 73(9):3059-63]을 참조한다.
알파바이러스 레플리콘은 복제를 위한 DNA 중간체를 포함하지 않으며, 레트로바이러스 벡터를 포함하여 일반적으로 사용되는 여러 다른 바이러스 벡터에 대한 보다 안전한 대안을 제공한다 ([Yoshioka et al., Cell Stem Cell. (2013) 13(2):246-54]; [Yoshioka and Dowdy, PLOS ONE (2017) 12:e0182018]). 알파바이러스, 및 특히, VEE는 체세포를 유도 만능 줄기 세포 (iPSC)로 재프로그래밍할 때 외인성 전사 인자를 코딩하는 유전자를 운반하는 벡터로 연구되었다. 그러나, 이러한 접근법은 섬유모세포 및 혈액 파생 내피 세포 (BOEC)에서만 시도되었다. 두 세포 유형 모두 임상적으로 특히 매력적이지는 않다. 자가 섬유모세포는 침습적이고 고통스러운 환자의 피부 천자로부터 수득된다. BOEC는 비록 말초 혈액으로부터 유래되기는 하지만, 희귀한 세포이며, 확립하는 데 힘이 들고, 시간이 많이 소요되는 프로세스가 요구된다.
통합 없는 또 다른 재프로그래밍 접근 방법은 에피솜 DNA 플라스미드 벡터를 이용한다. 웬(Wen) 등은 말초 혈액 단핵 세포를 iPSC로 재프로그래밍하는 데 상기 접근법을 사용하였다 (Stem Cell Rep. (2016) 6:873-84). 그러나, 상기 접근법의 경우, 다중 벡터를 통해 도입되는 다양한 재프로그래밍 인자 수준을 주의 깊게 보정하여야 한다.
센다이 바이러스 벡터 또한 재프로그래밍 인자를 코딩하는 유전자를 운반하는데 사용되어 왔다. 센다이 벡터는 (-) 가닥 파라믹소바이러스 (Paramyxovirus)이고; 벡터는 비리온으로 패키징되어야 한다. 이러한 접근법은 더 복잡하다. 이는 세포주를 패키징하는 것을 포함하고, 벡터 생성물에 외래성 오염 인자가 유입될 수 있다.
따라서, 말초 혈액 세포로부터 iPSC를 수득하기 위한 효율적이고, 안전한 접근법이 요구되고 있다.
본 개시내용은 조혈 계통의 출발 세포로부터 유도 만능 줄기 세포 (iPSC) 집단을 수득하는 방법을 제공한다. 본 방법은 BCL-xL, 및 OCT 패밀리 단백질, KLF 패밀리 단백질, MYC 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물을 출발 세포에 도입하는 단계, 및 출발 세포를 배양하여 BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자를 발현시켜 출발 세포 및 그의 자손이 iPSC로 재프로그래밍되도록 유도하는 단계를 포함한다.
한 측면에서, 본 개시내용은 BCL-xL, 및 Oct 패밀리 단백질, KLF 패밀리 단백질, Myc 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물로 형질감염된 조혈 계통의 출발 세포로부터 수득된 유도 만능 줄기 세포 (iPSC) 집단을 제공한다.
출발 세포는 예를 들어, 인간 기원의 조혈 줄기 세포, 적혈구 전구 세포, 림프양 전구 세포, 말초 혈액 단핵 세포, T 림프구, B 림프구, 대식세포, 단핵구, 호중구, 호산구, 또는 수지상 세포일 수 있다. 일부 실시양태에서, 출발 세포는 에리트로포이에틴 (EPO), 줄기 세포 인자 (SCF), 및 IL-3의 존재하에 임의적으로 5 내지 10일, 또는 6 내지 7일 동안 말초 혈액 단핵 세포 (PBMC)를 배양함으로써 수득된 적혈구 전구 세포이다. 추가 실시양태에서, PBMC는 0.5-5 IU/ml EPO, 50-200 ng/mL SCF, 및 1-10 ng/mL IL-3의 존재하에 배양된다.
일부 실시양태에서, RNA 발현 구축물은 전기천공을 통해 출발 세포에 도입된다. 추가 실시양태에서, 출발 세포는 전기천공 전에 B18R 단백질과 함께 배양된다.
또 다른 측면에서, 본 개시내용은 BCL-xL, 및 OCT 패밀리 단백질, KLF 패밀리 단백질, MYC 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물을 제공한다.
본원의 알파바이러스 RNA 발현 구축물에 대한 코딩 서열을 포함하는 DNA 벡터, 및 본원의 알파바이러스 RNA 발현 구축물 또는 DNA 벡터를 포함하는 숙주 세포 (예컨대, 인간 세포) 또한 제공한다.
일부 실시양태에서, 알파바이러스 RNA 발현 구축물은 자기 복제성이고, 구축물이 자기 복제되도록 하는 데 충분한 하나 이상의 비구조 단백질에 대한 유전자(예컨대, nsP1, nsP2, nsP3, 및 nsP4 유전자)를 포함한다. 추가 실시양태에서, 알파바이러스 RNA 발현 구축물은 베네수엘라 말 뇌염 바이러스 (VEEV) RNA 발현 구축물이고, VEEV nsP1, nsP2, nsP3, 및 nsP4 유전자를 포함한다. 특정 실시양태에서, VEEV RNA 발현 구축물은 야생형 VEEV 게놈의 상응하는 영역(들)으로부터 하나 이상의 (예컨대, 2개 이상, 3개 이상, 4개 이상, 5개 이상, 또는 6개 이상) 돌연변이를 함유한다.
일부 실시양태에서, OCT 패밀리 단백질은 OCT4 (예컨대, 인간 OCT4)이고/거나; KLF 패밀리 단백질은 KLF4 (예컨대, 인간 KLF4)이고/거나; SOX 패밀리 단백질은 SOX2 (예컨대, 인간 SOX2)이고/거나; LIN28 단백질은 LIN28B (예컨대, 인간 LIN28B)이고/거나; MYC 패밀리 단백질은 c-MYC (예컨대, 인간 c-MYC)이다.
특정 실시양태에서, 서열식별번호(SEQ ID NO:) 1 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 BCL-xL 단백질; 서열식별번호 3 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 OCT4 단백질; 서열식별번호 5 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 KLF4 단백질; 서열식별번호 7 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 SOX2 단백질; 및/또는 서열식별번호 9 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 c-MYC 단백질.
일부 실시양태에서, BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자에 대한 코딩 서열은 2A 펩티드 또는 내부 리보솜 진입 부위 (IRES)에 대한 코딩 서열에 의해 이격되어 있다. 일부 실시양태에서, BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자에 대한 코딩 서열은 공통 프로모터 (예컨대, 26S 프로모터)의 전사 제어하에 있다.
일부 실시양태에서, 본원의 알파바이러스 RNA 발현 구축물은 OCT 패밀리 단백질, SOX 패밀리 단백질, BCL-xL, 및 MYC 패밀리 단백질, 및 임의적으로 KLF 패밀리 단백질의 발현을 지시한다.
또 다른 측면에서, 본 개시내용은 분화 촉진제의 존재하에 본원에서 수득된 iPSC를 배양하는 단계를 포함하는, 시험관내에서 분화된 세포를 수득하는 방법을 제공한다. 또한, iPSC로부터의 분화에 의해 수득된 분화된 세포도 제공한다. 일부 실시양태에서, 본원에서 수득된 분화된 세포는 임의적으로 T 세포, 키메라 항원 수용체 (CAR)를 발현하는 T 세포, 억제 T 세포, 골수성 세포, 수지상 세포, 및 면역억제 대식세포로부터 선택되는 인간 면역 세포; 임의적으로 도파민성 뉴런, 미세아교세포, 희소돌기아교세포, 성상세포, 피질 뉴런, 척수 또는 안구운동 뉴런, 장 뉴런, 기원판 유래 세포, 슈반 세포, 및 삼차신경 또는 감각 뉴런으로부터 선택되는 인간 신경계 세포; 임의적으로 심근세포, 내피 세포, 및 결절 세포로부터 선택되는 인간 심혈관계 세포; 임의적으로 간세포, 담관세포, 및 췌장 베타 세포로부터 선택되는 인간 대사계 세포, 또는 임의적으로 망막 색소 상피 세포, 광수용체 추상 세포, 광수용체 간상 세포, 양극 세포, 신경절 세포로부터 선택되는 인간 안구계 세포, 면역 세포, 신경 세포, 심혈관 세포, 또는 대사계 세포이다. 특정 실시양태에서, 분화된 세포는 외배엽 계통의 것 (예컨대, 뉴런)이다. 다른 특정 실시양태에서, 분화된 세포는 중배엽 계통의 것 (예컨대, 심근세포)이다.
본 개시내용은 또한 본원에서 수득된 분화된 세포 및 제약상 허용되는 담체를 포함하는 제약 조성물을 제공한다. 본 개시내용은 또한 치료를 필요로 하는 환자에게 상기 제약 조성물을 투여하는 단계를 포함하는, 치료를 필요로 하는 환자를 치료하는 방법; 치료를 필요로 하는 환자를 치료하기 위한 의약의 제조를 위한 분화된 세포의 용도; 및 치료를 필요로 하는 환자 치료에서 사용하기 위한 분화된 세포 또는 제약 조성물도 제공한다.
본 발명의 다른 특징, 목적, 및 이점은 하기 상세한 설명에서 명백해진다. 그러나, 상세한 설명은 본 발명의 실시양태 및 측면을 나타내지만, 제한이 아닌 단지 예시로서 제공되는 것임을 이해하여야 한다. 본 발명의 범주 내에서의 다양한 변경 및 수정이 상세한 설명으로부터 관련 기술분야의 통상의 기술자에게 명백해질 것이다.
도 1은 7개의 예시적인 VEEV RNA 재프로그래밍 구축물 (OKS-iBM, OKS-iGM, OKS-iG, OSB, OS-iB, OS-iM, 및 OS-iBM)을 도시한 개략도이다. nsP1-4: 비구조 단백질에 대한 코딩 서열. OCT4: 팔량체 결합 전사 인자 4에 대한 코딩 서열. KLF4: 크루펠 유사 인자 4에 대한 코딩 서열. SOX2: SRY-박스 전사 인자 2에 대한 코딩 서열. BCL-xL: B 세포 림프종 - 초대형. GLIS1: Glis 패밀리 아연 핑거 1에 대한 코딩 서열. IRES: 내부 리보솜 진입 부위. 별표 표시된 회색 막대: 2A 펩티드에 대한 코딩 서열. AAA: 폴리-A 서열.
도 2a-d는 4개의 각 VEE-EP-iPSC 세포주, VEE-EP-iPSC-1, -2, -3 및 -4에서의 만능성 연관 마커의 발현을 보여주는 유세포 분석 그래프이다. 적혈구 전구 (EP) 세포를 각각 OKS-iBM, OKS-iGM, OKS-iG, 및 에피솜 대조군으로 전기천공하여 VEE-EP-iPSC-1, -2, -3 및 -4를 생성하였다. EBNA OriP 에피솜 대조군은 재프로그래밍 인자 OCT4, SOX2, KLF4, L-MYC, LIN28, 및 p53 우성 음성을 함유하였다 (Epi5™ 에피솜 iPSC 재프로그래밍 키트(Epi5™ Episomal iPSC Reprogramming Kit), 써모 피셔(Thermo Fisher); 문헌 [Okita et al., Nat Meth . (2011) 8:409-12]). 8 계대에서 배양된 세포에 대해 유세포 분석법을 수행하였다.
도 3은 먼저 신경외배엽 계통을 유도한 후, 이어서, 전구체를 신경 운명으로 성숙화시킴으로써 4개의 VEE-EP-iPSC 세포주가 16일 동안 분화되었다는 것을 보여주는 막대 그래프이다. TH+FOXA2+ 도파민 뉴런은 유세포 분석법으로 정량화하였다. TH: 티로신 히드록실라제. FOXA2: 포크헤드 박스 단백질 A2.
도 4는 4개의 VEE-EP-iPSC 세포주가 WNT 신호전달의 단계 특이적 조정을 통해 7일 동안 심장 계통으로 분화되었다는 것을 보여주는 막대 그래프이다. 유세포 분석법에 의해 심근세포를 심장 트로포닌 (cTNT)에 대해 정량화하였다.
도 5a 및 5b는 EP의 VEE RNA 재프로그래밍 효율을 보여주는 것이다. EP를 도 1에 도시된 재프로그래밍 구축물로 전기천공시켰다. 도 2에 대하여 상기 기술된 것과 같은 에피솜 대조군을 사용하였다. 도 5a는 TRA-1-60 염색의 전체 웰 이미징을 보여주는 사진 패널이다. 도 5b는 TRA-1-60+ 콜로니를 정량화한 그래프이다.
도 6a는 야생형 VEEV RNA 게놈 서열의 뉴클레오티드 서열 (서열식별번호 14)을 보여주는 것이다 (단, 서열에서 T는 RNA인 경우에는 U이다).
도 6b는 재조합 VEEV RNA 발현 벡터의 뉴클레오티드 서열 (서열식별번호 15)을 보여주는 것이다 (단, 서열에서 T는 RNA인 경우에는 U이다). 본 서열은 야생형 서열과 비교하여 6개의 돌연변이 (C352G, A1564G, C1567A, T1570C, C1647A, 및 C3917T)를 포함한다. 클로닝 부위는 별표로 표시되어 있다.
도 2a-d는 4개의 각 VEE-EP-iPSC 세포주, VEE-EP-iPSC-1, -2, -3 및 -4에서의 만능성 연관 마커의 발현을 보여주는 유세포 분석 그래프이다. 적혈구 전구 (EP) 세포를 각각 OKS-iBM, OKS-iGM, OKS-iG, 및 에피솜 대조군으로 전기천공하여 VEE-EP-iPSC-1, -2, -3 및 -4를 생성하였다. EBNA OriP 에피솜 대조군은 재프로그래밍 인자 OCT4, SOX2, KLF4, L-MYC, LIN28, 및 p53 우성 음성을 함유하였다 (Epi5™ 에피솜 iPSC 재프로그래밍 키트(Epi5™ Episomal iPSC Reprogramming Kit), 써모 피셔(Thermo Fisher); 문헌 [Okita et al., Nat Meth . (2011) 8:409-12]). 8 계대에서 배양된 세포에 대해 유세포 분석법을 수행하였다.
도 3은 먼저 신경외배엽 계통을 유도한 후, 이어서, 전구체를 신경 운명으로 성숙화시킴으로써 4개의 VEE-EP-iPSC 세포주가 16일 동안 분화되었다는 것을 보여주는 막대 그래프이다. TH+FOXA2+ 도파민 뉴런은 유세포 분석법으로 정량화하였다. TH: 티로신 히드록실라제. FOXA2: 포크헤드 박스 단백질 A2.
도 4는 4개의 VEE-EP-iPSC 세포주가 WNT 신호전달의 단계 특이적 조정을 통해 7일 동안 심장 계통으로 분화되었다는 것을 보여주는 막대 그래프이다. 유세포 분석법에 의해 심근세포를 심장 트로포닌 (cTNT)에 대해 정량화하였다.
도 5a 및 5b는 EP의 VEE RNA 재프로그래밍 효율을 보여주는 것이다. EP를 도 1에 도시된 재프로그래밍 구축물로 전기천공시켰다. 도 2에 대하여 상기 기술된 것과 같은 에피솜 대조군을 사용하였다. 도 5a는 TRA-1-60 염색의 전체 웰 이미징을 보여주는 사진 패널이다. 도 5b는 TRA-1-60+ 콜로니를 정량화한 그래프이다.
도 6a는 야생형 VEEV RNA 게놈 서열의 뉴클레오티드 서열 (서열식별번호 14)을 보여주는 것이다 (단, 서열에서 T는 RNA인 경우에는 U이다).
도 6b는 재조합 VEEV RNA 발현 벡터의 뉴클레오티드 서열 (서열식별번호 15)을 보여주는 것이다 (단, 서열에서 T는 RNA인 경우에는 U이다). 본 서열은 야생형 서열과 비교하여 6개의 돌연변이 (C352G, A1564G, C1567A, T1570C, C1647A, 및 C3917T)를 포함한다. 클로닝 부위는 별표로 표시되어 있다.
본 개시내용은 혈액 유래 세포 (예컨대, 적혈구 전구체)를 유도 만능 줄기 세포 (iPSC)로 재프로그래밍하기 위한 개선된 방법을 기술한다. 본 방법은 재프로그래밍 인자 BCL-xL, 및 하나 이상의 (예컨대, 1, 2, 3, 4, 5, 6, 7, 또는 8개 모두) 추가의 재프로그래밍 인자 (예컨대, OCT 패밀리 구성원, KLF 패밀리 구성원, SOX 패밀리 구성원, MYC 단백질, NANOG 단백질, GLIS 패밀리 구성원, LIN28 단백질, 및 p53 우성 음성)를 코딩하는 알파바이러스 (예컨대, VEEV) RNA 발현 벡터 (즉, 또는 발현 구축물)의 사용을 포함한다. 알파바이러스 RNA 발현 구축물은 본원에 기술된 개선된 방법을 통해 혈액 세포에 도입될 수 있다. 형질감염된 세포는 3주 미만의 기간 내에 수확 가능한 iPSC로 발생한다.
말초 혈액은 체세포를 iPSC로 재프로그래밍하기 위한 것으로서, 쉽게 이용할 수 있는 세포 공급원이다. 따라서, 본 방법은 iPSC 생성 효율을 크게 향상시킨다. 숙주 세포에 통합되지 않는 RNA 기반 발현 벡터를 사용하기 때문에, 본 방법에 의해 수득된 iPSC는 레트로바이러스 벡터를 사용한 이전 방법에 의해 수득된 것보다 더 안전한 임상 프로파일을 가진다.
I.
재프로그래밍 인자를 발현시키기 위한
알파바이러스
RNA 구축물
본 개시내용의 알파바이러스 RNA 발현 구축물은 자기 복제 RNA 레플리콘이다. 자기 복제 RNA 레플리콘 또는 구축물은 숙주 세포에서 그 자신의 복제를 지시할 수 있도록 비구조 단백질 유전자를 발현하는 RNA 분자를 지칭한다. 이는 5' 및 3' 알파바이러스 복제 인식 서열, RNA 복제 및 전사에 필수적인 알파바이러스 비구조 단백질에 대한 코딩 서열 (예컨대, VEE nsP1, nsP2, nsP3, 및 nsP4) 및 폴리아데닐화 신호 서열을 포함할 수 있다. 이는 추가로 재프로그래밍 인자를 코딩하는 것과 같은 이종성 RNA 서열의 발현을 지시하는 하나 이상의 요소 (예컨대, IRES 서열, 코어 또는 미니-프로모터 등)를 함유할 수 있다.
일부 실시양태에서, 알파바이러스 RNA 구축물은 (i) 복제에 필요한 VEEV 비구조 단백질에 대한 유전자, (ii) 5' 및 3' 바이러스 복제 인식 서열, (iii) 관심 재프로그래밍 인자를 발현시키기 위한, 발현 카세트(들), 예컨대, 폴리시스트론 발현 카세트; 및 (iv) 폴리아데닐화 테일을 포함하는 VEEV RNA 레플리콘이다. 문헌 [Yoshioka 2013 and 2017, 상기 문헌 동일]; 및 WO 2013/177133, 및 미국 특허 10,793,833, 10,370,646, 및 9,862,930 또한 참조한다. 레플리콘에는 VEEV 구조 단백질 유전자기 결여되어 있을 수 있다. 자기 복제 VEE RNA 구축물은 제한된 수의 세포 분열 동안 형질감염된 세포 내부에서 복제될 수 있다. 분해에 의한 RNA 구축물 손실 타이밍은 배양 배지에서 B18R을 제거함으로써 추가로 조절될 수 있다.
예시적인 VEEV RNA 구축물은 BCL-xL 및 다른 재프로그래밍 인자를 발현한다. 재프로그래밍 인자는 체세포에서 과다발현될 때, 세포가 분화 상태에서 만능 상태로 이행되도록 유도하는 단백질이다. 본원에서 사용되는 재프로그래밍 인자는 원하는 생물학적 효과를 유지하는 인간 단백질 또는 그의 변형된 버전일 수 있다.
A.
BCL
-
xL
인간 BCL-xL은 BCL2L1 유전자에 의해 코딩된다. 예시적인 인간 BCL-xL 아미노산 서열은 UniProt 수탁 번호 Q07817에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 BCL-xL 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 1과 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다. 두 아미노산 서열의 (또는 두 핵산 서열의) 동일성(%)은 예컨대, 디폴트 파라미터를 사용하여 BLAST® (미국 국립 의학 도서관의 국립 생명공학 정보 센터(U.S. National Library of Medicine's National Center for Biotechnology Information) 웹사이트에서 이용가능)에 의해 수득할 수 있다. 일부 실시양태에서, 비교 목적을 위해 정렬된 참조 서열의 길이는 참조 서열의 적어도 30% (예컨대, 적어도 40, 50, 60, 70, 80, 또는 90%)이다.
특정 실시양태에서, 본원의 구축물에 의해 발현된 BCL-xL 단백질은 하기 서열을 갖고, 여기서 박스내 잔기는 프로세싱 후 2A 자기 절단 펩티드로부터의 잔여물이다 (상이한 자기 절단 펩티드는 다른 잔여물을 남길 수 있거나, 또는 잔여물이 없을 수도 있다):
B. OCT
패밀리
구성원
예시적인 VEEV 구축물은 Oct 패밀리 단백질 (예컨대, OCT1, OCT2, OCT4, OCT6, OCT7, OCT8, OCT9, 및 OCT11)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 8,278,104 및 WO 2013/177133을 참조한다. 인간 OCT4는 POU5F1 유전자에 의해 코딩된다. 예시적인 인간 OCT4 아미노산 서열은 UniProt 수탁 번호 Q01860에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 OCT4 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 3과 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다.
특정 실시양태에서, 본원의 구축물에 의해 발현된 OCT4 단백질은 하기 서열을 갖고, 여기서 박스내 잔기는 프로세싱 후 2A 자기 절단 펩티드로부터의 잔여물이다 (상이한 자기 절단 펩티드는 다른 잔여물을 남길 수 있거나, 또는 잔여물이 없을 수도 있다):
C.
KLF
패밀리
구성원
예시적인 VEEV 구축물은 KLF 패밀리 단백질 (예컨대, KLF1, KLF2, KLF3, KLF4, KLF5, KLF6, KLF7, KLF8, KLF9, KLF10, KLF11, KLF12, KLF13, KLF14, KLF15, KLF16, 및 KLF17)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 8,278,104 및 WO 2013/177133을 참조한다. 인간 KLF4는 KLF4 유전자에 의해 코딩된다. 예시적인 인간 KLF4 아미노산 서열은 UniProt 수탁 번호 O43474에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 KLF4 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 5와 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다. 일부 실시양태에서, ESSRB가 KLF 단백질 대신 사용될 수 있다. 일부 실시양태에서, KLF4 단백질은 서열식별번호 5의 이소폼이고, 하기 제시된 서열식별번호 6의 아미노산 잔기 2-471을 포함한다.
특정 실시양태에서, 본원의 구축물에 의해 발현된 KLF4 단백질은 하기 서열을 갖고, 여기서 박스내 잔기는 프로세싱 후 2A 자기 절단 펩티드로부터의 잔여물이다 (상이한 자기 절단 펩티드는 다른 잔여물을 남길 수 있거나, 또는 잔여물이 없을 수도 있다):
D. SOX
패밀리
구성원
예시적인 VEEV 구축물은 SOX 패밀리 단백질 (예컨대, SOX1, SOX2, SOX3, SOX4, SOX5, SOX6, SOX7, SOX8, SOX9, SXO10, SOX11, SOX12, SOX13, SOX14, SOX15, SOX17, SOX18, SOX21, 및 SOX30)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 8,278,104 및 WO 2013/177133을 참조한다. 인간 SOX2는 SOX2 유전자에 의해 코딩된다. 예시적인 인간 SOX2 아미노산 서열은 UniProt 수탁 번호 P48431에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 SOX2 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 7과 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다.
특정 실시양태에서, 본원의 구축물에 의해 발현된 SOX2 단백질은 하기 서열을 갖고, 여기서 박스내 잔기는 프로세싱 후 2A 자기 절단 펩티드로부터의 잔여물이다 (상이한 자기 절단 펩티드는 다른 잔여물을 남길 수 있거나, 또는 잔여물이 없을 수도 있다):
E.
MYC
패밀리
구성원
예시적인 VEEV 구축물은 MYC 패밀리 단백질 (예컨대, c-MYC, n-MYC, 및 l-MYC)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 8,278,104를 참조한다. 인간 c-MYC는 MYC 유전자에 의해 코딩된다. 예시적인 인간 c-MYC 아미노산 서열은 UniProt 수탁 번호 P01106에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 c-MYC 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 9와 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다. 일부 실시양태에서, 형질전환 활성이 감소된 MYC 변이체는 c-MYC 대신 사용될 수 있다. 예컨대, 미국 특허 9,005,967을 참조한다.
특정 실시양태에서, 본원의 구축물에 의해 발현된 c-MYC 단백질은 하기 서열을 갖고, 여기서 박스내 잔기는 프로세싱 후 2A 자기 절단 펩티드로부터의 잔여물이다 (상이한 자기 절단 펩티드는 다른 잔여물을 남길 수 있거나, 또는 잔여물이 없을 수도 있다):
F.
GLIS
패밀리
구성원
예시적인 VEEV 구축물은 GLIS 패밀리 단백질 (예컨대, GLIS1, GLIS2, 및 GLIS3)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 8,951,801을 참조한다. 인간 GLIS1은 GLIS1 유전자에 의해 코딩된다. 예시적인 인간 GLIS1 아미노산 서열은 UniProt 수탁 번호 Q8NBF1에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 단백질의 기능적 유사체, 즉, 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 GLIS1 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 11과 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다.
G.
NANOG
본 VEEV 구축물은 NANOG에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 9,506,039를 참조한다. 인간 NANOG는 NANOG 유전자에 의해 코딩된다. 예시적인 인간 NANOG 아미노산 서열은 UniProt 수탁 번호 Q9H9S0에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 서열의 기능적 유사체, 즉, 상기 단백질의 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 NANOG 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 12와 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다.
H.
LIN28
단백질
본 VEEV 구축물은 LIN28 단백질 (예컨대, LIN28A 또는 LIN28B)에 대한 코딩 서열을 포함할 수 있다. 예컨대, 미국 특허 9,506,039를 참조한다. 인간 LIN28B는 LIN28B 유전자에 의해 코딩된다. 예시적인 인간 LIN28B 아미노산 서열은 UniProt 수탁 번호 A0A1B0GVD3에서 확인할 수 있고, 하기 아미노산 서열을 갖는다:
상기 서열의 기능적 유사체, 즉, 상기 단백질의 동일하거나, 또는 실질적으로 동일한 생물학적 기능을 갖는 (예컨대, 단백질의 전사 인자 기능을 70% 이상, 80% 이상, 90% 이상, 95% 이상 또는 98% 이상 유지하는) 분자는 LIN28B 단백질로서 본 개시내용에 포함된다. 예를 들어, 기능적 유사체는 상기 단백질의 이소폼 또는 변이체, 예컨대, 추가의 아미노산 잔기와 함께, 또는 그러한 잔기 없이 상기 단백질의 일부를 함유하고/거나, 상기 단백질 대비 돌연변이를 함유하는 것일 수 있다. 일부 실시양태에서, 기능적 유사체는 서열식별번호 13과 적어도 90, 95, 98, 또는 99%의 서열 동일성을 갖는다.
본원에 기술된 재프로그래밍 인자의 예시적인 기능적 유사체는 예컨대, 문헌 [Yang et al., Asian J Andrology (2015) 17:394-402] (상기 문헌의 개시내용은 그 전문이 본원에서 참조로 포함된다)에 기술되어 있다.
I. RNA 발현 구축물
일부 실시양태에서, 재프로그래밍 인자의 코딩 서열은, 각각이 각각은 그 자신의 프로모터 (예컨대, 26S 프로모터) 및 다른 전사 조절 요소를 갖는 하나 이상의 발현 카세트에 포함될 수 있다.
일부 실시양태에서, 재프로그래밍 인자의 코딩 서열은 공통 프로모터 (예컨대, 26S 또는 SP6 프로모터)로부터 전사되도록 폴리시스트론 발현 카세트 중 프레임 내에 배치될 수 있다. 이들 코딩 서열은 번역 스키핑 서열 (즉, 프레임내의 자기 절단 펩티드에 대한 코딩 서열)에 의해 이격될 수 있고, 이로써, 폴리시스트론 카세트로부터의 mRNA 전사체의 번역으로 인해 별개의 단백질이 생성될 것이다. 자기 절단 펩티드는 번역 중에 리보솜 스키핑을 유발한다. 자기 절단 펩티드의 예는 2A 펩티드이며, 이는 전형적인 길이가 18-22개의 아미노산 길이인 바이러스 유래 펩티드이다. 2A 펩티드는 T2A, P2A, E2A, F2A 및 PQR을 포함한다 (Lo et al., Cell Reports (2015) 13:2634-2644). 예를 들어, P2A는 19개의 아미노산으로 이루어진 펩티드이며; 절단 후 P2A의 아미노산 잔기 몇 개가 상류 유전자에 남고, 프롤린은 2번째 유전자의 시작 부분에 남다. 재프로그래밍 인자에 대한 코딩 서열은 또한 mRNA 중 내부 리보솜 진입 부위 (IRES)에 의해 이격될 수도 있다. IRES는 또한 일반 RNA 전사체로부터 별개의 폴리펩티드를 번역하는 것을 허용한다. 프로세싱된 폴리펩티드에 남아 있는 2A 잔기는 폴리펩티드의 기능성에 영향을 미치지 않다.
예를 들어, 알파바이러스 RNA 구축물은 5'에서 3'으로 하기를 포함할 수 있다: [알파바이러스 5' UTR] - [알파바이러스 RNA 레플리카제에 대한 유전자] - [프로모터] - [재프로그래밍 인자 1 코딩 서열] - [2A 펩티드 코딩 서열] - [재프로그래밍 인자 2 코딩 서열] - [2A 펩티드 코딩 서열] - [재프로그래밍 인자 3 코딩 서열] - [IRES 또는 코어 프로모터] - [재프로그래밍 인자 4 코딩 서열] - [2A 펩티드 코딩 서열] - [재프로그래밍 인자 5] - [임의적 선별가능한 마커] - [알파바이러스 3' UTR 및 폴리A 테일]. 폴리A 테일의 길이는 달라질 수 있고 (예컨대, 10 내지 200개 초과의 아데노신 길이), 재프로그래밍 인자의 순서는 RNA 구축물의 재프로그래밍 기능에 영향을 미치지 않으면서 변경될 수 있다. 폴리시스트론 재프로그래밍 인자 발현 카세트에 대한 프로모터는 예를 들어, 26S 내부 프로모터일 수 있다. 일부 실시양태에서, 알파바이러스 RNA 구축물은 VEEV RNA 구축물이고, 그의 레플리카제에 대한 유전자는 VEEV RNA 레플리카제 1, 2, 3, 및 4이다.
일부 실시양태에서, 알파바이러스 (예컨대, VEEV) RNA 구축물은 임의적으로 5'에서 3'으로 nsP1, nsP2, nsP3, 및 nsP4에 대한 코딩 서열, 및 재프로그래밍 인자의 조합, 예컨대, (i) OCT4, KLF4, SOX2, BCL-xL, 및 c-Myc, (ii) OCT4, SOX2, BCL-xL, 및 c-MYC, 또는 (ii) OCT4, SOX2, 및 BCL-xL의 발현을 위한 폴리시스트론 발현 카세트를 포함하는, 도 1에 제시된 바와 같은 구조를 가질 수 있다. 폴리시스트론 발현 카세트에서, 발현 카세트는 26S 프로모터의 전사 제어하에 있을 수 있고/거나, 재프로그래밍 인자에 대한 코딩 서열은 IRES 서열 또는 자기 절단 2A 펩티드에 대한 코딩 서열에 의해 이격될 수 있다 (예를 들어, IRES 위치의 비제한적인 예는 도 1에 제시되어 있다).
알파바이러스 (예컨대, VEEV) RNA 구축물은 DNA 주형 (예컨대, DNA 플라스미드 구축물)으로부터 생산될 수 있다. 예를 들어, RNA 구축물은 SP6 (또는 T7) 시험관내 전사 키트를 사용하여 DNA 주형으로부터 전사될 수 있다.
임의의 VEEV 균주를 사용하여 본 RNA 구축물에 대한 백본을 제공할 수 있다. 예를 들어, VEEV의 TC-83 균주를 사용할 수 있다. 이 균주는 nsP2에 P773S 돌연변이를 포함하고 있으며, 결과적으로 형질도입된 세포에 대해 감소된 세포변성 효과를 미친다. RNA 복제 및 발현을 개선시키고/거나, RNA 게놈에 대한 면역 반응을 약화시키기 위해 ns 단백질 중 하나 이상에 다른 또는 추가의 돌연변이가 도입될 수 있다. 예를 들어, VEEV RNA 발현 구축물은 도 6b에 제시된 돌연변이 중 하나 이상 (예컨대, 2, 3, 4, 5, 또는 6개 모두)을 포함할 수 있다. 추가로, 재프로그래밍 자기 복제 RNA 구축물에 대한 백본을 제공하기 위해 VEEV 대신 다른 알파바이러스 또한 사용될 수 있다. RNA 구축물의 기반이 될 수 있는 다른 알파바이러스의 비제한적 예로는 동부 말 뇌염 바이러스 (EEEV), 에버글레이즈 바이러스(Everglades virus), 무캄보 바이러스(Mucambo virus), 픽수나 바이러스(Pixuna virus) 및 서부 말 뇌염 바이러스 (WEEV), 신드비스 바이러스, 셈리키 포레스트 바이러스(Semliki Forest virus), 미들버그 바이러스(Middelburg virus), 치쿤군야 바이러스, 오뇽뇽 바이러스, 로스 리버 바이러스, 바마 포레스트 바이러스(Barmah Forest virus), 게타 바이러스(Getah virus), 사기야마 바이러스(Sagiyama virus), 베바루 바이러스(Bebaru virus), 마야로 바이러스(Mayaro virus), 우나 바이러스(Una virus), 아우라 바이러스(Aura virus), 와타로아 바이러스(Whataroa virus), 바방키 바이러스(Babanki virus), 키질라가흐 바이러스(Kyzylagach virus), 하이랜드 J 바이러스(Highlands J virus), 포트 모건 바이러스(Fort Morgan virus), 엔두무 바이러스(Ndumu virus) 및 버기 크릭 바이러스(Buggy Creek virus)가 있다. RNA 구축물은 1 초과의 알파바이러스로부터의 서열을 함유할 수 있다.
II.
RNA 구축물을 사용한 조혈 계통의 세포로부터의
iPSC
생성
본 방법은 혈액 세포가 유도 만능 줄기 세포가 되도록 효율적으로 재프로그래밍 (또는 "탈분화"로도 명명)한다.
본원에서 사용되는 바와 같이, 용어 "만능" 또는 "만능성"은 자기 재생하고, 3개의 배엽층: 내배엽, 중배엽 또는 외배엽 중 어느 것으로 분화하는 세포의 능력을 지칭한다. "만능 줄기 세포" 또는 "PSC"는 예를 들어 배반포의 내부 세포 매스로부터 유래되거나, 체세포 핵 이식에 의해 유래된 배아 줄기 세포, 및 비만능 세포로부터 유래된 iPSC를 포함한다.
용어 "유도 만능 줄기 세포" 또는 "iPSC"는 비만능 세포, 예컨대, 성체 체세포, 부분 분화된 세포 또는 최종 분화된 세포, 예컨대, 섬유모세포, 조혈 계통의 세포, 근세포, 뉴런, 표피 세포 등에 하나 이상의 재프로그래밍 인자를 도입하거나, 또는 그와 접촉시킴으로써 상기 세포로부터 인공적으로 제조되는 만능 줄기 세포 타입을 지칭한다.
PSC 유도를 위한 출발 세포 집단은 세포 요법이 필요한 환자로부터의 또는 건강한 공여자로부터의 혈액 (예컨대, 말초 혈액)으로부터 수득될 수 있다. 말초 혈액 단핵 세포 (PMBC)는 종래 방법으로 단리시킨 후, 이어서, 추가로 분별 및/또는 농축시켜 세포의 서브세트, 예컨대, T 림프구, B 림프구, 단핵구, 자연 살해 세포, 호중구, 호산구, 수지상 세포 및 각종 조혈 전구 세포, 예컨대, 적혈구 전구체, 림프양 전구체, 및 골수성 전구체를 수득할 수 있다.
일부 실시양태에서, PMBC를 에리트로포이에틴 (EPO), 줄기 세포 인자 (SCF) 및 IL-3으로 보충된 기본 배지 (예컨대, 스템스판(StemSpan)™ SFEM II 배지; 스템셀 테크놀러지즈(StemCell Technologies)) 중에서 일정 기간 (예컨대, 3-10일, 예컨대, 6, 7, 또는 8일) 동안 배양하여 적혈구 전구 세포에 대해 농축된 세포 집단을 수득한다. 배양 배지를 예를 들어, 0.5 내지 5 (예컨대, 1, 2, 3, 또는 4) IU/mL EPO, 50 내지 200 (예컨대, 75, 100, 125, 150, 또는 175) ng/mL SCF, 및 1 내지 10 (예컨대, 2, 3, 4, 5, 6, 7, 8, 또는 9) ng/mL IL-3으로 보충할 수 있다. 적혈구 전구체 증식을 촉진시키는 다른 인자, 예를 들어, 재조합 인간 인슐린, 철 포화 인간 트랜스페린, 질산제2철, 히드로코르티손 또한 사용될 수 있다. 예컨대, 문헌 [Neildez-Nguyen et al., Nat Biotechnol . (2002) 20:467-72]; 및 [Filippone et al., PLoS One (2010) 5(3):e9496]을 참조한다. 적혈구 전구체는 예컨대, 적혈구 전구체 마커, 예컨대, CD71 및 CD36에 결합하는 시약 (예컨대, 항체)을 사용하여 형광 또는 자기 활성화 세포 분류에 의해 세포 배양물로부터 추가로 단리될 수 있다.
일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 100 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 100 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 150 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 150 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 200 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 1 IU/ml EPO, 약 200 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 100 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 100 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 150 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 150 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 200 ng/mL SCF, 및 약 5 ng/mL IL-3의 존재하에서 배양한다. 일부 실시양태에서, PBMC를 3 IU/ml EPO, 약 200 ng/mL SCF, 및 약 10 ng/mL IL-3의 존재하에서 배양한다. 상기 실시양태에서, 배양은 6, 7, 또는 8일 동안 수행될 수 있다.
혈액 세포의 다른 서브세트 또한 세포 배양을 거쳐 분별 및/또는 농축에 의해 수득될 수 있다. T 림프구의 경우, CD3, 및 B 세포의 경우 CD19 및 CD20과 같이 혈액 세포의 특정 서브세트에 대한 마커는 널리 공지되어 있다.
본 RNA 구축물은 미세주입, 전기천공, 바이올리스틱 입자 전달, 리포펙션, 양이온 중합체, 및 인산칼슘 침전을 비롯한 다수의 기술에 의해 체세포 집단 내로 도입될 수 있다. 일부 실시양태에서, 본 RNA 구축물은 전기천공을 통해 체세포 (예컨대, 조혈 전구 세포 및 림프구) 내로 도입된다. 알파바이러스를 벡터로 사용하는 것은 인터페론 (IFN)에 의한 선천적 면역 반응에 의해 억제될 수 있는 것으로 알려져 있지만, 예컨대, B18R 또는 B19R과 같은 타입 I IFN 억제제를 사용하여 세포 항바이러스 반응을 억제함으로써 세포에서 원하는 레플리콘 활성을 가능하게 할 수 있다. 일부 실시양태에서, 알파바이러스 (예컨대, VEEV) 전달 및 후속 복제를 촉진시키고/거나, 형질감염된 세포에서 세포 인터페론 반응을 억제시키기 위해 전기천공 전에 세포를 B18R 단백질로 처리할 수 있다. 전기천공된 세포는 B18R의 존재하에서 2-3주 동안 배양될 수 있고, 이 기간 동안 iPSC가 출현하고, 이를 수확할 수 있다. iPSC는 예컨대, TRA-1-60, NANOG, SSEA3, 및 SSEA4와 같은 마커에 의해 검출될 수 있다. 일부 실시양태에서, 예컨대, Opti-MEM® (써모 피셔)과 같은 배양 배지는 전기천공 후 세포의 생존을 촉진시키기 위한 전기천공 세포 현탁 완충제로 사용될 수 있다. 일부 실시양태에서, RNA 구축물은 알파바이러스 비리온으로 패키징될 수 있고, 비리온은 재프로그래밍되는 세포에의 형질도입에 사용된다.
iPSC를 유지하는 방법은 관련 기술분야에 널리 공지되어 있고, 상기 방법 중 다수는 배아 줄기 세포를 유지하는 방법과 유사하다. 예컨대, 문헌 [Thomson et al., Science (1998) 282(5391):1145-7]; [Hovatta et al., Human Reprod . (2003) 18(7):1404-09]; [Ludwig et al., Nature Methods (2006) 3:637-46]; [Kennedy et al., Blood (2007) 109:2679-87]; [Chen et al., Nature Methods (2011) 8:424-9]; 및 [Wang et al., Stem Cell Res. (2013) 11(3):1103-16]을 참조한다. iPSC 또한 사용 전에 냉동보존될 수 있다.
III.
iPSC의
표적 세포 유형으로의 분화
iPSC는 다수의 상이한 질환을 앓고 있는 환자의 재생 의학을 위해 전달될 수 있는 다수의 특정 세포 유형을 잠재적으로 생성할 수 있는 출발점이다. iPSC 맥락에서 분화는 iPSC부터 출발하여 세포 특이적 프로토콜을 사용한 계통 특정화 프로세스이다. 본 방법에 의해 수득된 iPSC는 내배엽, 외배엽 및 중배엽 계통의 세포를 비롯한, 세포 요법을 위한 관심 세포 유형으로 분화될 수 있다. 일부 실시양태에서, iPSC를 먼저 관심 세포 유형으로의 분화 전에 (예컨대, 환자에서 결함이 있는 기능적 단백질을 생산하기 위해, 치료 단백질을 생산하기 위해, 자살 스위치를 포함하기 위해, 또는 면역 검출을 회피하여 동종 이계 적용을 지원하기 위해) 유전자 조작할 수 있다. iPSC의 다양한 계통의 세포로의 분화 및 그의 확장을 유도하는 방법은 관련 기술분야에 널리 공지되어 있다. 분화된 세포 유형의 비제한적인 예는 하기 기술되어 있다.
A. 면역 세포
임의적으로 유전자 변형된 iPSC는 면역 세포, 예컨대, 림프양 세포 (예컨대, T 세포, B 세포, 및 NK 세포), 골수성 세포 (예컨대, 과립구, 단핵구/대식세포, 및 조직 상주 대식세포, 예컨대, 미세아교세포), 및 수지상 세포 (예컨대, 골수성 수지상 세포 및 형질세포성 수지상 세포)로 분화될 수 있다. 일부 실시양태에서, 유전자 변형된 세포는 키메라 항원 수용체 (CAR)를 발현하는 T 세포 또는 CAR T 세포이다. 유전자 변형된 면역 세포는 또한 면역조절 트랜스진, 예컨대, HLA-G 또는 HLA-E를 발현할 수 있다.
예를 들어, PSC의 수지상 세포로의 분화를 유도하는 방법은 문헌 [Slukvin et al., J Imm . (2006) 176:2924-32]; [Su et al., Clin Cancer Res. (2008) 14(19):6207-17]; 및 [Tseng et al., Regen Med . (2009) 4(4):513-26]에 기술되어 있다. PSC를 조혈 전구 세포, 골수성 계통의 세포, 및 T 림프구로 유도하는 방법은 예컨대, 문헌 [Kennedy et al., Cell Rep. (2012) 2:1722-35]에 기술되어 있다. PSC를 대식세포로 유도하는 방법은 문헌 [van Wilgenburg et al., PLoS One (2013) 8(8):e71098]에 기술되어 있다.
면역 세포, 예컨대, 면역억제 면역 세포 (예컨대, 조절성 T 세포 및 면역억제 대식세포)는 제한 없이, 류마티스 관절염, 다발성 경화증, 만성 림프구성 갑상선염, 인슐린 의존성 당뇨병, 중증 근무력증, 만성 궤양성 결장염, 궤양성 결장염, 크론병, 염증성 장 질환, 굿파스처 증후군, 전신 홍반성 루푸스, 전신 혈관염, 경피증, 자기면역 용혈성 빈혈, 자기면역 갑상선 질환을 비롯한, 자가면역 질환을 앓는 환자 내로 이식될 수 있다. 면역 세포 기반 요법은 예컨대, 섬유증과 같은 이식 관련 증상의 치료를 포함하여 이식시 이식 거부를 치료하는 데에도 사용될 수 있다.
B. 신경 세포
임의적으로 유전자 변형된 iPSC는 제한 없이, 임의의 특정 뉴런 서브타입 (예컨대, 도파민성 뉴런, 장 뉴런, 개재뉴런, 및 피질 뉴런)과 관계없이 뉴런 및 뉴런 전구 세포; 임의의 특정 신경교 서브타입 (예컨대, 희소돌기아교세포, 성상세포, 전용 희소돌기아교세포 전구 세포, 및 성상세포 및 희소돌기아교세포를 생성할 수 있는 이분화능 신경교 전구체)과 관계없이 신경교 세포 및 신경교 전구 세포; 및 미세아교세포 및 미세아교세포 전구 세포를 포함하는 신경 세포로 분화될 수 있다. 척수 또는 안구운동 뉴런, 장 뉴런, 기원판 유래 세포, 슈반 세포, 및 삼차신경 또는 감각 뉴런 또한 고려된다.
신경 세포는 제한 없이, 신경퇴행성 질환을 앓고 있는 환자를 비롯한 환자 내로 이식될 수 있다. 신경퇴행성 질환의 예로는 그 중에서도 특히 파킨슨병, 알츠하이머병, 치매, 간질, 루이체 증후군, 헌팅턴병, 척수성 근위축증, 프리드라이히 운동실조증(Hurler syndrome), 근위축성 측삭 경화증, 바텐병(Batten disease), 다계통 위축증, 백질이영양증, 횡단 척수염, 시신경척수염, 리소솜 축적병 (예컨대, 헐러 증후군(Hurler syndrome), 파브리병(Fabry disease), 고셔병(Gaucher disease), 슬라이 증후군(Sly syndrome), GM1 및 GM2 강글리오시드증, 헌터 증후군(Hunter syndrome), 니만-픽병(Niemann-Pick disease), 산필리포 증후군(Sanfilippo syndrome)), 타우병증이 있다.
이러한 다수의 질환의 경우, iPSC는 먼저 이중 SMAD 억제를 통해 원시 신경 세포 운명을 채택하도록 지시될 수 있다 (Chambers et al., Nat Biotechnol. (2009) 27(3):275-80). 원시 신경 세포는 전방 특징을 채택하는 바, 추가 신호가 없으면 전방/전뇌 피질 세포를 제공하게 될 것이다. 후방화 신호를 차단하여 더 후방 특징이 있는 배양물을 생성할 수 있는 측분비 신호를 방지할 수 있다 (예를 들어, XAV939는 WNT를 차단할 수 있고, SU5402는 FGF 신호를 차단할 수 있다). 등쪽 피질 뉴런은 SHH 활성화를 차단함으로써 생성될 수 있는 반면, 배쪽 피질 뉴런은 SHH 활성화를 통해 생성될 수 있다. 예컨대, 세로토닌성 뉴런 또는 척수 운동 뉴런과 같은 더 후방 세포 유형은 FGF 및/또는 WNT 신호의 추가를 통해 배양물을 후방화함으로써 생성될 수 있다. 일부 세포 유형의 경우, 레티노산 (또 다른 후방화제)을 첨가하여 배양물을 배면화할 수 있다. 신경교 세포 유형의 생산은 일반적으로 FGF2 및/또는 EGF 함유 배지에서의 확장 배양 전 원시 신경 세포의 동일한 패턴화를 따를 수 있다. PNS 세포 유형은 동일한 일반 원칙을 따를 수 있지만, 분화 프로세스에서 조기에 적시의 WNT 신호하에 이루어질 수 있다.
신경 세포는 손상된 해당 조직에 배치된 캐뉼라를 통해 환자 내로 도입될 수 있다. 세포 제제를 지지 배지에 넣고, 제제를 정확하게 전달할 수 있는 시린지 또는 피펫 유사 장치에 로딩할 수 있다. 이어서, 일반적으로 전달을 정확하게 표적화하는 정위 방법을 사용하여 캐뉼라를 환자의 신경계에 배치할 수 있다. 이어서, 세포는 적합한 속도로 조직으로 방출될 수 있다.
C. 심혈관 세포
임의적으로 유전자 변형된 iPSC는 심혈관계 세포, 예컨대, 특정 심근세포 서브타입 (예컨대, 심실 또는 심방)을 비롯한 심근세포, 심장 섬유모세포, 심장 평활근 세포, 심장 외심막 세포, 심장 내심막 세포, 심장 내피 세포, 푸르키니에 섬유(Purkinje fiber), 및 결절 및 페이스메이커 세포로 분화될 수 있다. 예를 들어, 문헌 [Kattman et al., Cell Stem Cell (2011) 8(2):228-40]; [Lian et al., PNAS (2012) 109:e1848-57]; [Lee et al., Cell Stem Cell (2017) 21:179-94]에 제시된 바와 같이, 및 WO 2016/131137, WO 2018/098597, 및 미국 특허 9,453,201에 제시된 바와 같이 iPSC를 심근세포로 분화시키는 방법이 다수 존재한다. 관련 기술분야의 임의의 적합한 방법을 본원의 방법과 함께 사용하여 PSC 유래 심근세포를 수득할 수 있다.
일부 실시양태에서, iPSC는 하나 이상의 심장 분화 배지 중에서 인큐베이션된다. 예를 들어, 배지는 다양한 농도의 골형성 단백질 (BMP, 예컨대, BMP4) 및 액티빈 (예컨대, 액티빈 A)을 함유할 수 있다. 원하는 심근세포 분화를 달성하는 데 필요한 최적의 농도를 결정하기 위해 분화 인자 농도의 적정을 수행할 수 있다.
일부 실시양태에서, 분화된 심근세포는 심장 트로포닌 T (cTnT), 및/또는 미오신 경쇄 2v (MLC2v) 중 하나 이상의 것을 발현한다. 일부 실시양태에서, 미성숙 심근세포는 트로포닌 T, 심장 트로포닌 I, 알파 액티닌 및/또는 베타-미오신 중쇄 중 하나 이상의 것을 발현한다.
D.
대사계의
세포
임의적으로 유전자 변형된 iPSC는 인간 대사계와 관련된 세포로 분화될 수 있다. 예를 들어, 세포는 위장계 세포 (예컨대, 간세포, 담관세포 및 췌장 베타 세포), 조혈계 세포, 및 중추 신경계 세포 (예컨대, 뇌하수체 호르몬 방출 세포)일 수 있다. 예를 들어, 뇌하수체 호르몬 방출 세포를 생성하기 위해, SHH/FGF8 및 FGF10을 첨가하기 전에 iPSC를 (액티빈 신호전달을 차단하는) BMP4 및 SB431542와 함께 배양한 후; 이어서, 세포에 FGF8 또는 BMP (또는 그 둘 모두) 전에 연장된 기간 동안 오직 SHH/FGF8 및 FGF10만을 가하여 세포가 특정 호르몬 방출 세포가 되도록 유도한다. 예컨대, 문헌 [Zimmer et al., Stem Cell Reports (2016) 6:858-72]를 참조한다.
E.
안구계의
세포
임의적으로 유전자 변형된 iPSC는 안구계의 세포로 분화될 수 있다. 예를 들어, 세포는 망막 전구 세포, 망막 색소 상피 (RPE) 전구 세포, RPE 세포, 신경 망막 전구 세포, 광수용체 전구 세포, 광수용체 세포, 양극 세포, 수평 세포, 신경절 세포, 아마크린 세포, 뮐러 신경교, 추상 세포, 또는 간상 세포일 수 있다. iPSC를 RPE 세포로 분화시키는 방법은 예컨대, WO 2017/044483에 기술되어 있다. RPE 세포를 단리시키는 방법은 예컨대, WO 2017/044488에 기술되어 있다. iPSC를 신경 망막 전구 세포로 분화시키는 방법은 WO 2019/204817에 기술되어 있다. 망막 전구 세포 및 RPE 세포를 확인 및 단리시키는 방법은 예컨대, WO 2011/028524에 기술되어 있다.
IV.
제약 조성물 및 용도
본원에 기술된 iPSC 유래 세포는 상기 세포 및 제약상 허용되는 담체를 함유하는 제약 조성물로 제공될 수 있다. 제약상 허용되는 담체는 임의적으로 동물 유래 성분을 전혀 함유하지 않는 세포 배양 배지일 수 있다. 보관 및 운송을 위해, 세포는 < -70℃에서 (예컨대, 드라이아이스 상에서 또는 액체 질소 중에서) 냉동보존될 수 있다. 사용 전, 세포를 해동하고, 관심 세포 유형을 지지하는 멸균 세포 배지 중에 희석할 수 있다.
세포는 환자에게 전신적으로 (예컨대, 정맥 주사 또는 주입을 통해) 또는 국소적으로 (예컨대, 국소 조직, 예컨대, 심장, 뇌 및 손상된 조직 부위에 직접 주사를 통해) 투여될 수 있다. 제한 없이, 관상동맥내 투여, 심근내 투여, 경심내막 투여 또는 두개내 투여를 비롯한, 환자의 조직 또는 기관 내로 세포를 투여하는 다양한 방법이 관련 기술분야에 공지되어 있다.
iPSC 유래 세포는 치료 유효 개수로 환자에게 투여된다. 본원에서 사용되는 바와 같이, "치료 유효"라는 용어는 질환, 장애, 및/또는 병태를 앓고 있거나, 이에 걸리기 쉬운 인간 대상체에게 투여되었을 때, 질환, 장애, 및/또는 병태를 치료, 예방하고/거나, 그의 증상(들)의 발병 또는 진행을 지연시키는 데 충분한 세포수 또는 제약 조성물의 양을 지칭한다. 치료 유효량은 전형적으로 적어도 1 단위 용량을 포함하는 투여 요법을 통해 투여된다는 것이 관련 기술분야의 통상의 기술자라면 이해할 수 있을 것이다.
본원에서 달리 정의되지 않는 한, 본 개시내용과 관련하여 사용되는 과학 및 기술 용어는 관련 기술분야의 통상의 기술자가 일반적으로 이해하는 의미를 가져야 한다. 본원에 기술된 것과 유사하거나 등가인 방법 및 물질 또한 본 개시내용의 실시 또는 테스팅에 사용될 수 있지만, 예시적인 방법 및 물질은 하기 기술된다. 상충하는 경우, 정의를 포함하는 본 명세서가 우선할 것이다. 일반적으로, 본원에 기술된 세포 및 조직 배양, 분자 생물학, 면역학, 미생물학, 유전학, 분석 화학, 합성 유기 화학, 의료 및 제약 화학, 단백질 및 핵산 화학 및 하이브리드화와 관련하여 사용되는 명명법, 및 그의 기술은 관련 기술분야에서 널리 공지되어 있고, 일반적으로 사용되는 것이다. 효소 반응 및 정제 기술은 관련 기술분야에서 일반적으로 수행되거나, 또는 본원에 기술된 바와 같이 제조업체의 사양에 따라 수행된다. 추가로, 문맥상 달리 요구되지 않는 한, 단수 용어는 복수를 포함하고, 복수 용어는 단수를 포함하여야 한다. 본 명세서 및 실시양태 전역에 걸쳐, "갖는다(have)" 및 "포함하다(comprise)"라는 단어 또는 예컨대, "갖는다(has)," "갖는," "포함하다(comprises)" 또는 "포함하는"과 같은 어미 변형은 언급된 정수 또는 정수 그룹의 포함을 의미하지만, 임의의 다른 정수 또는 정수 그룹의 배제를 암시하는 것은 아님을 이해할 것이다. 본원에서 언급된 모든 공개문헌 및 다른 참고문헌은 그 전문이 본원에서 참조로 포함된다. 본원에서 다수의 문헌이 인용되었지만, 이러한 인용은 이들 문헌 중 어느 것이 관련 기술분야의 일반 지식의 일부를 형성한다는 것을 인정하는 것으로 간주되지 않는다.
본 발명을 더 잘 이해할 수 있도록 하기 실시예를 설명한다. 이들 실시예는 단지 예시를 위한 것이며, 어느 방식으로든 본 발명의 범주를 제한하는 것으로 해석되지 않아야 한다.
실시예
실시예
1:
VEEV
디자인 및 RNA 합성
본 실시예는 폴리시스트론 VEEV RNA 구축물의 디자인 및 그의 합성을 기술한다. 하기 연구에서 사용된 구축물은 문헌 [Yoshioka, 2013, 상기 문헌 동일]에 기술된 VEEV RNA 백본 서열에 기반하였다. 상기 백본 서열은 VEEV의 4개의 비구조 단백질을 코딩한다. 재프로그래밍 핵 인자의 다양한 조합을 발현하도록 서브게놈 서열을 변형시켰다 (도 1). RNA 구축물 OKS-iBM은 OCT4, KLF4, SOX2, BCL-xL, 및 c-MYC를 코딩한다. RNA 구축물 OKS-iGM은 OCT4, KLF4, SOX2, GLIS1, 및 c-MYC를 코딩한다. RNA 구축물 OKS-iG는 OCT4, KLF4, SOX2, 및 GLIS1을 코딩한다. RNA 구축물 OSB는 OCT4, SOX2, 및 BCL-xL을 코딩한다. RNA 구축물 OS-iB는 OCT4, SOX2, 및 BCL-xL을 코딩한다. RNA 구축물 OS-iM은 OCT4, SOX2, 및 c-MYC를 코딩한다. RNA 구축물 OS-iBM은 OCT4, SOX2, BCL-xL 및 c-MYC를 코딩한다. 명칭에 "i"가 포함되어 있는 구축물은 SOX2 코딩 서열의 바로 하류에 IRES 서열을 함유한다. 다양한 재프로그래밍 인자에 대한 코딩 서열은 자기 절단 2A 펩티드에 대한 코딩 서열에 의해, 또는 IRES에 의해 이격되어 있고, 이로써, 모든 재프로그래밍 인자는 동일한 프로모터로부터 발현된다 ([Wen et al., Stem Cell Rep. (2016) 6:873-84]; [Su et al., PLoS ONE (2013) 8:e64496]).
VEEV RNA 구축물은 그의 각 DNA 플라스미드 주형으로부터 효소적으로 합성하였다. RNA 합성을 최적화하기 위해, AG 캡 아날로그 기술(AG Cap analog technology) (클린캡(CleanCap), 트리링크(Trilink))을 사용하여 5' 캡핑된 RNA를 생성하였다.
재조합 VEEV 구축물의 백본 서열이 도 6b (서열식별번호 15)에 제시되어 있고, 여기서 발현 카세트에 삽입 부위는 서열에서 별표로 표시되어 있다.
실시예
2: 적혈구 전구체의
iPSC로의
재프로그래밍
본 실시예는 적혈구 전구 세포를 iPSC로 재프로그래밍하기 위한 예시적인 프로토콜을 기술한다. 적혈구 전구체 (EP)에 대해 농축된 세포 집단을 수득하기 위해, 인간 공여자로부터의 PBMC를 해동시킨 후, 약 3 IU/mL EPO, 약 100 ng/mL SCF, 및 약 5 ng/mL IL-3으로 보충된 배지 ("EP 배지") 중에서 5 내지 10일 (예컨대, 6일) 동안 배양하였다. 상기 성장 인자는 EP 증식을 지원한다. 예컨대, 문헌 [Wang et al., Clin Hemorheol Microcirc. (2007) 37(4):291-9]를 참조한다. 대안적으로, 문헌 [Wen et al. (Stem Cell Reports (2016) 6:873-84)]에 기술된 바와 같이, PBMC를 예컨대, 100 ng/ml 줄기 세포 인자 (페프로테크(Peprotech); 300-07), 10 ng/ml 인터루킨-3 (페프로테크; AF-200-03), 2 U/ml 에리트로포이에틴 (페프로테크; 100-64), 20 ng/ml 인슐린 성장 인자-1 (페프로테크; 100-11), 1 mM 덱사메타손 (시그마((Sigma)); D4902), 및 0.2 mM 1-티오글리세롤 (시그마; M6145)로 보충된 스템라인® II 조혈 줄기 세포 확장 배지(Stemline® II Hematopoietic Stem Cell Expansion Medium) (시그마; S0192)를 포함하는 배양 배지 중에서 배양할 수 있다.
더욱 구체적으로, 조직 배양물로 처리된 플레이트 중 세포 밀도가 2-3x106 세포/mL를 달성하도록 해동된 PBMC를 EP 배지 중에 시딩하였다. 시딩 후 배지의 약 ¼ 내지 ¾을 밤새 (예컨대, 16-24 hr) 교체하였다. 2일째, 세포를 새 베쓸 (초저 부착성, 비조직 배양 처리)로 옮기고, 매일 25-75% 배지 교체를 수행하였다. 5일째, 추가 EP 배지를 첨가하여 세포를 2배로 희석시켰다. 6일째 (또는 7일째), 배양 배지의 절반을 교체하고, 유세포 분석법으로 EP 세포 샘플을 CD71 및 CD36에 대한 이중 양성에 대해 평가하였다.
이어서, CD71+CD36+ EP 세포를 20 min 동안 인터페론 억제자 (예컨대, 재조합 B18R 단백질)와 함께 인큐베이션시켰다. 세포를 원심분리하고, DPBS로 세척한 후, 이어서, Opti-MEM™ (써모 피셔 사이언티픽(Thermo Fisher Scientific)) 중에 약 2x107 세포/mL로 재현탁시켰다. 매 120 ㎕의 전기천공 반응마다 4 ㎍의 VEEV 재프로그래밍 RNA를 냉각된 1.5 mL 마이크로튜브로 옮겼다. 이어서, 세포를 전기천공하고, B18R이 보충된 EP 배지 중에 플레이팅하고, 2일 동안 유가식으로 배양하였다. 플레이트를 예컨대, 비트로넥틴 또는 라미닌과 같은 기질로 코팅하였다. 형질감염 후 3 내지 6일, 세포에 B18R로 보충된 에센셜 7(Essential 7) 또는 ReproTeSR™ 배지를 유가식으로 공급하였다. 형질감염 후 7일째를 시작으로 배양물을 매일 완전 배지로 교체하였다. iPSC 콜로니가 형질감염 후 약 10일째에 출현하였고, 15-20일째 사이에 선택할 준비가 되었다. 선택된 콜로니를 B18R 부재하에서 확장시킨 후, 이어서, 냉동보존하였다 (본원에서 VEE-EP-iPSC로 지칭).
도 2a-c에 제시된 바와 같이, 3개의 VEEV RNA 구축물로 생성된 VEE-EP-iPSC는 미분화 만능 세포와 연관된 핵 (NANOG) 및 표면 마커 (TRA-1-60, SSEA-3 및 SSEA-4)를 발현하였다. 상기 세포는 또한 정상적인 핵형을 가지고 있었다.
차세대 시퀀싱에 의해 VEE-EP-iPSC를 프로파일링하여 500개 초과의 암 연관 유전자에서의 유전자 변이체 획득을 평가하였다. VEE-EP-iPSC 세포주에서 500개 초과의 유전자의 유전자 서열을 출발 공여자 PBMC 집단과 비교하였을 때, 서열 차이는 관찰되지 않았다. 상기 데이터는 재프로그래밍 동안 유전자 변이체는 획득되지 않았다는 것을 입증한다.
모든 VEE-EP-iPSC 세포주는 외배엽을 나타내는 TH+ 도파민성 뉴런으로 분화하는 능력을 입증하였다 (도 3). VEE-EP-iPSC를 도파민성 뉴런으로 분화시키기 위해, 본 발명자들은 먼저 0일째부터 7일째까지 TGF-β 및 BMP 신호전달을 차단하여 신경외배엽 계통으로 iPSC를 유도하였다. 본 발명자들은 동시에 0일째부터 7일째까지 재조합 C25II SHH 단백질을 통해 세포를 바닥판 운명으로 패턴화하고, 0일째부터 12일째까지 WNT 효능제 소분자 CHIR-99021을 이용하여 세포의 중뇌 식별을 미세 조정하였다. 상기 단계 후, 본 발명자들은 10일째부터 16일째까지는 신경영양 인자를 사용하고, 12일째부터 16일째까지는 소분자 γ-세크레타제 억제제 DAPT를 사용하여 전구체를 뉴런 운명으로 성숙화시켰다. 16일째에 분화된 세포를 시험관내에서 5일 동안 성숙시키고, 유세포 분석법으로 TH+FOXA2+ 도파민 뉴런을 정량화하였다.
모든 VEE-EP-iPSC 세포주는 또한 중배엽 분화를 나타내는 심장 트로포닌 (cTNT) 양성 심근세포로 분화할 수 있었다 (도 4). VEE-EP-iPSC 세포주는 WNT 효능제 CHIR-99021 및 WNT 길항제 엔도-IWR1의 사용을 통한 WNT 신호전달의 단계 특이적 조정하에 심장 계통으로 분화되었다. 유세포 분석법에 의해 심근세포를 심장 트로포닌 (cTNT) 염색에 대하여 정량화하였다.
상이한 전사 인자 조합을 함유하는 VEEV RNA 구축물의 재프로그래밍 효율을 측정하기 위해, 적혈구 전구체를 PBMC로부터 확장시키고, 재프로그래밍 RNA 구축물로 전기천공시켰다. TRA-1-60 양성 콜로니를 전기천공 후 17일째 (OKS-iBM 및 에피솜 구축물의 경우) 또는 25일째 (OKS-iGM 및 OKS-iG 구축물의 경우)에 정량화하였다. 재프로그래밍 효율은 PSC 마커 TRA-1-60을 발현하는 세포 기반 콜로니의 개수로 측정하였다 (도 5a). VEEV RNA 매개 EP 재프로그래밍의 효율은 BCL-xL 코딩 서열이 VEEV 구축물에 포함되었을 때 유의하게 증가하였다 (도 5b). BCL-xL 코딩 서열을 포함한 VEEV OKS-iBM 구축물은 종래 재프로그래밍 인자 OCT4, SOX2, KLF4, L-MYC, LIN28, 및 p53 우성 음성을 포함하는 에피솜 기반 대조군보다 4배 더 높은 EP 재프로그래밍 효율을 입증하였다 (도 5b - iBM/ Epi5). OS-iBM 구축물은 또한 활성이었고, iPSC 콜로니를 형성하였다 (데이터는 제시되지 않음).
수(Su) 등 (2013, 상기 문헌 동일)이 제5 재프로그래밍 인자로서 BCL-xL을 포함하였을 때 (에피솜 OS+MK+B 조합), OS+MK (OCT4, SOX2, c-MYC 및 KLF4) 조합과 비교하였을 때, 재프로그래밍 효율이 대략 8배 증가한 것으로 관찰되었다. 그러나, 요시오카(Yoshioka) 및 다우디(Dowdy) (2017, 상기 문헌 동일)가 재프로그래밍 인자 조합을 평가하였을 때에는 4-인자 조합 (VEE-OKS-iM)과 비교하였을 때, GLIS1을 포함하는 것 (VEE-OKS-iGM)이 재프로그래밍 효율을 대략 20배만큼 부스팅하였다. 따라서, GLIS1을 BCL-xL로 대체하면 (VEE-OKS-iBM) 적혈구 전구체의 재프로그래밍 효율이 추가로 8배만큼 증가한 것은 예상치 못한 일이었다 (도 5b - iBM/iGM). 이러한 재프로그래밍 효율의 놀랍고도, 극적인 증가를 통해 효율이 낮은 적혈구 재프로그래밍은 강건한 접근법으로 전환된다. 더욱 강건한 효율은 상이한 혈액 샘플에 걸친 일관된 재프로그래밍을 허용하며, 임상 사용을 위한 EP 재프로그래밍의 실행가능성에 매우 중요하다.
실시예
3: T 세포의
iPSC로의
재프로그래밍
본 실시예는 T 림프구를 iPSC로 재프로그래밍하기 위한 프로토콜을 기술한다. 2명의 독립된 공여자로부터의 말초 혈액 (올셀즈(AllCells))의 음성 면역 선택 및 면역 표현형 분석을 통해 정제된 CD3+ T 세포 (범(pan)-T 세포)를 수득하였다. 두 공여자 모두, 그로부터의 범-T 세포를 해동시키고, CTS™ 글루타MAX™(CTS™ GlutaMAX™) 및 100 IU/mL IL-로 보충된 T 세포 완전 배지 중에 유지시켰다. 전기천공 전, 범-T 세포를 30 min 동안 0.2 ㎍/mL 재조합 B18R 단백질로 처리하여, 냉 포스페이트 완충처리된 염수로 세척하고, 재현탁시켰다.
세포를 전기천공시키고, 플레이팅하고, 0.2 ㎍/mL의 B18R, CTS™ 글루타MAX™, 및 100 IU/mL IL-2로 보충된 상기 T 세포 배지 중에서 24 hr 동안 초저 부착 플레이트 (코닝(Corning)) 상에서 인큐베이션시켰다. 이어서, 세포를 0.2 ㎍/mL B18R이 새로 보충된, LN521로 코팅된 조직 배양 플레이트 상에 재플레이팅하였다. 전기천공 후 3일째 내지 6일째 사이, 재프로그래밍 배양물에 스템핏 베이직03 배지(StemFit Basic03 medium) (아지노모토(Ajinomoto))를 유가식으로 공급하였고, 매일 0.2 ㎍/mL의 B18R을 보충해 주었다. 형질감염 후 7일째를 시작으로 배양물을 매일 B18R로 보충된 스템핏 베이직03 배지를 이용하여 완전 배지로 교체하였다. iPSC 콜로니가 형질감염 후 약 10일째에 출현하였고, 15-20일째 사이에 선택할 준비가 되었다. 선택된 콜로니를 B18R 부재하에서 스템핏 베이직03 배지를 이용하여 확장시킨 후, 이어서, 냉동보존하였다 (본원에서 VEE-T-iPSC로 지칭).
결론적으로, 본 발명자들은 적혈구 전구체 외에도 CD3+ T 세포가 VEE-OKS-iBM RNA를 사용한 전기천공에 의해 재프로그래밍될 수 있다는 것을 입증하였다. 상이한 두 공여자 T 세포 로트를 재프로그래밍하여 iPSC 세포주를 확립하였고, 여기서, 두 공여자의 평균 재프로그래밍 효율은 0.005%였다. 이는 형질감염된 세포 백만개당 50개의 콜로니를 생성하게 되는 바, 충분한 세포주 유도율이다.
서열 목록
본 개시내용의 예시적인 서열은 하기 표에서 제공된다 (SEQ: 서열식별번호 ).
<표 1>
SEQUENCE LISTING
<110> BLUEROCK THERAPEUTICS LP
<120> METHODS FOR OBTAINING INDUCED PLURIPOTENT STEM CELLS
<130> 025450.WO014
<140>
<141>
<150> 63/166,071
<151> 2021-03-25
<160> 15
<170> PatentIn version 3.5
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Ser Ala Ile Asn Gly Asn Pro Ser Trp His Leu Ala Asp Ser Pro Ala
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Val Asn Gly Ala Thr Gly His Ser Ser Ser Leu Asp Ala Arg Glu Val
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Ile Pro Met Ala Ala Val Lys Gln Ala Leu Arg Glu Ala Gly Asp Glu
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Phe Glu Leu Arg Tyr Arg Arg Ala Phe Ser Asp Leu Thr Ser Gln Leu
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His Ile Thr Pro Gly Thr Ala Tyr Gln Ser Phe Glu Gln Val Val Asn
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Glu Leu Phe Arg Asp Gly Val Asn Trp Gly Arg Ile Val Ala Phe Phe
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Ser Phe Gly Gly Ala Leu Cys Val Glu Ser Val Asp Lys Glu Met Gln
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Val Leu Val Ser Arg Ile Ala Ala Trp Met Ala Thr Tyr Leu Asn Asp
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His Leu Glu Pro Trp Ile Gln Glu Asn Gly Gly Trp Asp Thr Phe Val
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Glu Leu Tyr Gly Asn Asn Ala Ala Ala Glu Ser Arg Lys Gly Gln Glu
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Arg Phe Asn Arg Trp Phe Leu Thr Gly Met Thr Val Ala Gly Val Val
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Ser Ala Ile Asn Gly Asn Pro Ser Trp His Leu Ala Asp Ser Pro Ala
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Pro Pro Tyr Glu Phe Cys Gly Gly Met Ala Tyr Cys Gly Pro Gln Val
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Glu Pro Cys Thr Val Thr Pro Gly Ala Val Lys Leu Glu Lys Glu Lys
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Glu Leu Glu Gln Phe Ala Lys Leu Leu Lys Gln Lys Arg Ile Thr Leu
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Gly Tyr Thr Gln Ala Asp Val Gly Leu Thr Leu Gly Val Leu Phe Gly
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Lys Val Phe Ser Gln Thr Thr Ile Cys Arg Phe Glu Ala Leu Gln Leu
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Ser Phe Lys Asn Met Cys Lys Leu Arg Pro Leu Leu Gln Lys Trp Val
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Val Arg Gly Asn Leu Glu Asn Leu Phe Leu Gln Cys Pro Lys Pro Thr
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Phe Ser Gly Gly Pro Val Ser Phe Pro Leu Ala Pro Gly Pro His Phe
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Gly Thr Pro Gly Tyr Gly Ser Pro His Phe Thr Ala Leu Tyr Ser Ser
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Val Pro Phe Pro Glu Gly Glu Ala Phe Pro Pro Val Ser Val Thr Thr
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Leu Gly Ser Pro Met His Ser Asn
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<212> PRT
<213> Artificial Sequence
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Met Ala Gly His Leu Ala Ser Asp Phe Ala Phe Ser Pro Pro Pro Gly
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Gly Gly Gly Asp Gly Pro Gly Gly Pro Glu Pro Gly Trp Val Asp Pro
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Arg Thr Trp Leu Ser Phe Gln Gly Pro Pro Gly Gly Pro Gly Ile Gly
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Pro Gly Val Gly Pro Gly Ser Glu Val Trp Gly Ile Pro Pro Cys Pro
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Pro Pro Tyr Glu Phe Cys Gly Gly Met Ala Tyr Cys Gly Pro Gln Val
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Gly Val Gly Leu Val Pro Gln Gly Gly Leu Glu Thr Ser Gln Pro Glu
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Gly Glu Ala Gly Val Gly Val Glu Ser Asn Ser Asp Gly Ala Ser Pro
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Glu Pro Cys Thr Val Thr Pro Gly Ala Val Lys Leu Glu Lys Glu Lys
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Leu Glu Gln Asn Pro Glu Glu Ser Gln Asp Ile Lys Ala Leu Gln Lys
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Glu Leu Glu Gln Phe Ala Lys Leu Leu Lys Gln Lys Arg Ile Thr Leu
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Gly Tyr Thr Gln Ala Asp Val Gly Leu Thr Leu Gly Val Leu Phe Gly
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Ser Phe Lys Asn Met Cys Lys Leu Arg Pro Leu Leu Gln Lys Trp Val
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Thr Leu Val Gln Ala Arg Lys Arg Lys Arg Thr Ser Ile Glu Asn Arg
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Val Arg Gly Asn Leu Glu Asn Leu Phe Leu Gln Cys Pro Lys Pro Thr
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Leu Leu Thr Cys Gly Asp Val Glu Glu Asn Pro Gly
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Thr Leu Arg Gln Ala Gly Ala Pro Asn Asn Arg Trp Arg Glu Glu Leu
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Ser Ala Pro Pro Pro Thr Ala Pro Phe Asn Leu Ala Asp Ile Asn Asp
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Glu Tyr Gly Ser Pro Ser Val Ile Ser Val Ser Lys Gly Ser Pro Asp
245 250 255
Gly Ser His Pro Val Val Val Ala Pro Tyr Asn Gly Gly Pro Pro Arg
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Thr Cys Pro Lys Ile Lys Gln Glu Ala Val Ser Ser Cys Thr His Leu
275 280 285
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290 295 300
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305 310 315 320
Leu Glu Glu Val Leu Ser Ser Arg Asp Cys His Pro Ala Leu Pro Leu
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Pro Pro Gly Phe His Pro His Pro Gly Pro Asn Tyr Pro Ser Glu Leu
340 345 350
Met Pro Pro Gly Ser Cys Met Pro Glu Glu Pro Lys Pro Lys Arg Gly
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370 375 380
Ala Gly Cys Gly Lys Thr Tyr Thr Lys Ser Ser His Leu Lys Ala His
385 390 395 400
Arg Ser Asp His Leu Ala Leu His Met Lys Arg His Phe
405 410
<210> 6
<211> 493
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
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<400> 6
Pro Met Ala Val Ser Asp Ala Leu Leu Pro Ser Phe Ser Thr Phe Ala
1 5 10 15
Ser Gly Pro Ala Gly Arg Glu Lys Thr Leu Arg Gln Ala Gly Ala Pro
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Asn Asn Arg Trp Arg Glu Glu Leu Ser His Met Lys Arg Leu Pro Pro
35 40 45
Val Leu Pro Gly Arg Pro Tyr Asp Leu Ala Ala Ala Thr Val Ala Thr
50 55 60
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65 70 75 80
Ala Pro Leu Pro Arg Arg Glu Thr Glu Glu Phe Asn Asp Leu Leu Asp
85 90 95
Leu Asp Phe Ile Leu Ser Asn Ser Leu Thr His Pro Pro Glu Ser Val
100 105 110
Ala Ala Thr Val Ser Ser Ser Ala Ser Ala Ser Ser Ser Ser Ser Pro
115 120 125
Ser Ser Ser Gly Pro Ala Ser Ala Pro Ser Thr Cys Ser Phe Thr Tyr
130 135 140
Pro Ile Arg Ala Gly Asn Asp Pro Gly Val Ala Pro Gly Gly Thr Gly
145 150 155 160
Gly Gly Leu Leu Tyr Gly Arg Glu Ser Ala Pro Pro Pro Thr Ala Pro
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Phe Asn Leu Ala Asp Ile Asn Asp Val Ser Pro Ser Gly Gly Phe Val
180 185 190
Ala Glu Leu Leu Arg Pro Glu Leu Asp Pro Val Tyr Ile Pro Pro Gln
195 200 205
Gln Pro Gln Pro Pro Gly Gly Gly Leu Met Gly Lys Phe Val Leu Lys
210 215 220
Ala Ser Leu Ser Ala Pro Gly Ser Glu Tyr Gly Ser Pro Ser Val Ile
225 230 235 240
Ser Val Ser Lys Gly Ser Pro Asp Gly Ser His Pro Val Val Val Ala
245 250 255
Pro Tyr Asn Gly Gly Pro Pro Arg Thr Cys Pro Lys Ile Lys Gln Glu
260 265 270
Ala Val Ser Ser Cys Thr His Leu Gly Ala Gly Pro Pro Leu Ser Asn
275 280 285
Gly His Arg Pro Ala Ala His Asp Phe Pro Leu Gly Arg Gln Leu Pro
290 295 300
Ser Arg Thr Thr Pro Thr Leu Gly Leu Glu Glu Val Leu Ser Ser Arg
305 310 315 320
Asp Cys His Pro Ala Leu Pro Leu Pro Pro Gly Phe His Pro His Pro
325 330 335
Gly Pro Asn Tyr Pro Ser Phe Leu Pro Asp Gln Met Gln Pro Gln Val
340 345 350
Pro Pro Leu His Tyr Gln Glu Leu Met Pro Pro Gly Ser Cys Met Pro
355 360 365
Glu Glu Pro Lys Pro Lys Arg Gly Arg Arg Ser Trp Pro Arg Lys Arg
370 375 380
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385 390 395 400
Lys Ser Ser His Leu Lys Ala His Leu Arg Thr His Thr Gly Glu Lys
405 410 415
Pro Tyr His Cys Asp Trp Asp Gly Cys Gly Trp Lys Phe Ala Arg Ser
420 425 430
Asp Glu Leu Thr Arg His Tyr Arg Lys His Thr Gly His Arg Pro Phe
435 440 445
Gln Cys Gln Lys Cys Asp Arg Ala Phe Ser Arg Ser Asp His Leu Ala
450 455 460
Leu His Met Lys Arg His Phe Gly Ser Gly Gln Cys Thr Asn Tyr Ala
465 470 475 480
Leu Leu Lys Leu Ala Gly Asp Val Glu Ser Asn Pro Gly
485 490
<210> 7
<211> 317
<212> PRT
<213> Homo sapiens
<400> 7
Met Tyr Asn Met Met Glu Thr Glu Leu Lys Pro Pro Gly Pro Gln Gln
1 5 10 15
Thr Ser Gly Gly Gly Gly Gly Asn Ser Thr Ala Ala Ala Ala Gly Gly
20 25 30
Asn Gln Lys Asn Ser Pro Asp Arg Val Lys Arg Pro Met Asn Ala Phe
35 40 45
Met Val Trp Ser Arg Gly Gln Arg Arg Lys Met Ala Gln Glu Asn Pro
50 55 60
Lys Met His Asn Ser Glu Ile Ser Lys Arg Leu Gly Ala Glu Trp Lys
65 70 75 80
Leu Leu Ser Glu Thr Glu Lys Arg Pro Phe Ile Asp Glu Ala Lys Arg
85 90 95
Leu Arg Ala Leu His Met Lys Glu His Pro Asp Tyr Lys Tyr Arg Pro
100 105 110
Arg Arg Lys Thr Lys Thr Leu Met Lys Lys Asp Lys Tyr Thr Leu Pro
115 120 125
Gly Gly Leu Leu Ala Pro Gly Gly Asn Ser Met Ala Ser Gly Val Gly
130 135 140
Val Gly Ala Gly Leu Gly Ala Gly Val Asn Gln Arg Met Asp Ser Tyr
145 150 155 160
Ala His Met Asn Gly Trp Ser Asn Gly Ser Tyr Ser Met Met Gln Asp
165 170 175
Gln Leu Gly Tyr Pro Gln His Pro Gly Leu Asn Ala His Gly Ala Ala
180 185 190
Gln Met Gln Pro Met His Arg Tyr Asp Val Ser Ala Leu Gln Tyr Asn
195 200 205
Ser Met Thr Ser Ser Gln Thr Tyr Met Asn Gly Ser Pro Thr Tyr Ser
210 215 220
Met Ser Tyr Ser Gln Gln Gly Thr Pro Gly Met Ala Leu Gly Ser Met
225 230 235 240
Gly Ser Val Val Lys Ser Glu Ala Ser Ser Ser Pro Pro Val Val Thr
245 250 255
Ser Ser Ser His Ser Arg Ala Pro Cys Gln Ala Gly Asp Leu Arg Asp
260 265 270
Met Ile Ser Met Tyr Leu Pro Gly Ala Glu Val Pro Glu Pro Ala Ala
275 280 285
Pro Ser Arg Leu His Met Ser Gln His Tyr Gln Ser Gly Pro Val Pro
290 295 300
Gly Thr Ala Ile Asn Gly Thr Leu Pro Leu Ser His Met
305 310 315
<210> 8
<211> 318
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
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<400> 8
Pro Met Tyr Asn Met Met Glu Thr Glu Leu Lys Pro Pro Gly Pro Gln
1 5 10 15
Gln Thr Ser Gly Gly Gly Gly Gly Asn Ser Thr Ala Ala Ala Ala Gly
20 25 30
Gly Asn Gln Lys Asn Ser Pro Asp Arg Val Lys Arg Pro Met Asn Ala
35 40 45
Phe Met Val Trp Ser Arg Gly Gln Arg Arg Lys Met Ala Gln Glu Asn
50 55 60
Pro Lys Met His Asn Ser Glu Ile Ser Lys Arg Leu Gly Ala Glu Trp
65 70 75 80
Lys Leu Leu Ser Glu Thr Glu Lys Arg Pro Phe Ile Asp Glu Ala Lys
85 90 95
Arg Leu Arg Ala Leu His Met Lys Glu His Pro Asp Tyr Lys Tyr Arg
100 105 110
Pro Arg Arg Lys Thr Lys Thr Leu Met Lys Lys Asp Lys Tyr Thr Leu
115 120 125
Pro Gly Gly Leu Leu Ala Pro Gly Gly Asn Ser Met Ala Ser Gly Val
130 135 140
Gly Val Gly Ala Gly Leu Gly Ala Gly Val Asn Gln Arg Met Asp Ser
145 150 155 160
Tyr Ala His Met Asn Gly Trp Ser Asn Gly Ser Tyr Ser Met Met Gln
165 170 175
Asp Gln Leu Gly Tyr Pro Gln His Pro Gly Leu Asn Ala His Gly Ala
180 185 190
Ala Gln Met Gln Pro Met His Arg Tyr Asp Val Ser Ala Leu Gln Tyr
195 200 205
Asn Ser Met Thr Ser Ser Gln Thr Tyr Met Asn Gly Ser Pro Thr Tyr
210 215 220
Ser Met Ser Tyr Ser Gln Gln Gly Thr Pro Gly Met Ala Leu Gly Ser
225 230 235 240
Met Gly Ser Val Val Lys Ser Glu Ala Ser Ser Ser Pro Pro Val Val
245 250 255
Thr Ser Ser Ser His Ser Arg Ala Pro Cys Gln Ala Gly Asp Leu Arg
260 265 270
Asp Met Ile Ser Met Tyr Leu Pro Gly Ala Glu Val Pro Glu Pro Ala
275 280 285
Ala Pro Ser Arg Leu His Met Ser Gln His Tyr Gln Ser Gly Pro Val
290 295 300
Pro Gly Thr Ala Ile Asn Gly Thr Leu Pro Leu Ser His Met
305 310 315
<210> 9
<211> 439
<212> PRT
<213> Homo sapiens
<400> 9
Met Pro Leu Asn Val Ser Phe Thr Asn Arg Asn Tyr Asp Leu Asp Tyr
1 5 10 15
Asp Ser Val Gln Pro Tyr Phe Tyr Cys Asp Glu Glu Glu Asn Phe Tyr
20 25 30
Gln Gln Gln Gln Gln Ser Glu Leu Gln Pro Pro Ala Pro Ser Glu Asp
35 40 45
Ile Trp Lys Lys Phe Glu Leu Leu Pro Thr Pro Pro Leu Ser Pro Ser
50 55 60
Arg Arg Ser Gly Leu Cys Ser Pro Ser Tyr Val Ala Val Thr Pro Phe
65 70 75 80
Ser Leu Arg Gly Asp Asn Asp Gly Gly Gly Gly Ser Phe Ser Thr Ala
85 90 95
Asp Gln Leu Glu Met Val Thr Glu Leu Leu Gly Gly Asp Met Val Asn
100 105 110
Gln Ser Phe Ile Cys Asp Pro Asp Asp Glu Thr Phe Ile Lys Asn Ile
115 120 125
Ile Ile Gln Asp Cys Met Trp Ser Gly Phe Ser Ala Ala Ala Lys Leu
130 135 140
Val Ser Glu Lys Leu Ala Ser Tyr Gln Ala Ala Arg Lys Asp Ser Gly
145 150 155 160
Ser Pro Asn Pro Ala Arg Gly His Ser Val Cys Ser Thr Ser Ser Leu
165 170 175
Tyr Leu Gln Asp Leu Ser Ala Ala Ala Ser Glu Cys Ile Asp Pro Ser
180 185 190
Val Val Phe Pro Tyr Pro Leu Asn Asp Ser Ser Ser Pro Lys Ser Cys
195 200 205
Ala Ser Gln Asp Ser Ser Ala Phe Ser Pro Ser Ser Asp Ser Leu Leu
210 215 220
Ser Ser Thr Glu Ser Ser Pro Gln Gly Ser Pro Glu Pro Leu Val Leu
225 230 235 240
His Glu Glu Thr Pro Pro Thr Thr Ser Ser Asp Ser Glu Glu Glu Gln
245 250 255
Glu Asp Glu Glu Glu Ile Asp Val Val Ser Val Glu Lys Arg Gln Ala
260 265 270
Pro Gly Lys Arg Ser Glu Ser Gly Ser Pro Ser Ala Gly Gly His Ser
275 280 285
Lys Pro Pro His Ser Pro Leu Val Leu Lys Arg Cys His Val Ser Thr
290 295 300
His Gln His Asn Tyr Ala Ala Pro Pro Ser Thr Arg Lys Asp Tyr Pro
305 310 315 320
Ala Ala Lys Arg Val Lys Leu Asp Ser Val Arg Val Leu Arg Gln Ile
325 330 335
Ser Asn Asn Arg Lys Cys Thr Ser Pro Arg Ser Ser Asp Thr Glu Glu
340 345 350
Asn Val Lys Arg Arg Thr His Asn Val Leu Glu Arg Gln Arg Arg Asn
355 360 365
Glu Leu Lys Arg Ser Phe Phe Ala Leu Arg Asp Gln Ile Pro Glu Leu
370 375 380
Glu Asn Asn Glu Lys Ala Pro Lys Val Val Ile Leu Lys Lys Ala Thr
385 390 395 400
Ala Tyr Ile Leu Ser Val Gln Ala Glu Glu Gln Lys Leu Ile Ser Glu
405 410 415
Glu Asp Leu Leu Arg Lys Arg Arg Glu Gln Leu Lys His Lys Leu Glu
420 425 430
Gln Leu Arg Asn Ser Cys Ala
435
<210> 10
<211> 440
<212> PRT
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polypeptide"
<400> 10
Pro Met Pro Leu Asn Val Ser Phe Thr Asn Arg Asn Tyr Asp Leu Asp
1 5 10 15
Tyr Asp Ser Val Gln Pro Tyr Phe Tyr Cys Asp Glu Glu Glu Asn Phe
20 25 30
Tyr Gln Gln Gln Gln Gln Ser Glu Leu Gln Pro Pro Ala Pro Ser Glu
35 40 45
Asp Ile Trp Lys Lys Phe Glu Leu Leu Pro Thr Pro Pro Leu Ser Pro
50 55 60
Ser Arg Arg Ser Gly Leu Cys Ser Pro Ser Tyr Val Ala Val Thr Pro
65 70 75 80
Phe Ser Leu Arg Gly Asp Asn Asp Gly Gly Gly Gly Ser Phe Ser Thr
85 90 95
Ala Asp Gln Leu Glu Met Val Thr Glu Leu Leu Gly Gly Asp Met Val
100 105 110
Asn Gln Ser Phe Ile Cys Asp Pro Asp Asp Glu Thr Phe Ile Lys Asn
115 120 125
Ile Ile Ile Gln Asp Cys Met Trp Ser Gly Phe Ser Ala Ala Ala Lys
130 135 140
Leu Val Ser Glu Lys Leu Ala Ser Tyr Gln Ala Ala Arg Lys Asp Ser
145 150 155 160
Gly Ser Pro Asn Pro Ala Arg Gly His Ser Val Cys Ser Thr Ser Ser
165 170 175
Leu Tyr Leu Gln Asp Leu Ser Ala Ala Ala Ser Glu Cys Ile Asp Pro
180 185 190
Ser Val Val Phe Pro Tyr Pro Leu Asn Asp Ser Ser Ser Pro Lys Ser
195 200 205
Cys Ala Ser Gln Asp Ser Ser Ala Phe Ser Pro Ser Ser Asp Ser Leu
210 215 220
Leu Ser Ser Thr Glu Ser Ser Pro Gln Gly Ser Pro Glu Pro Leu Val
225 230 235 240
Leu His Glu Glu Thr Pro Pro Thr Thr Ser Ser Asp Ser Glu Glu Glu
245 250 255
Gln Glu Asp Glu Glu Glu Ile Asp Val Val Ser Val Glu Lys Arg Gln
260 265 270
Ala Pro Gly Lys Arg Ser Glu Ser Gly Ser Pro Ser Ala Gly Gly His
275 280 285
Ser Lys Pro Pro His Ser Pro Leu Val Leu Lys Arg Cys His Val Ser
290 295 300
Thr His Gln His Asn Tyr Ala Ala Pro Pro Ser Thr Arg Lys Asp Tyr
305 310 315 320
Pro Ala Ala Lys Arg Val Lys Leu Asp Ser Val Arg Val Leu Arg Gln
325 330 335
Ile Ser Asn Asn Arg Lys Cys Thr Ser Pro Arg Ser Ser Asp Thr Glu
340 345 350
Glu Asn Val Lys Arg Arg Thr His Asn Val Leu Glu Arg Gln Arg Arg
355 360 365
Asn Glu Leu Lys Arg Ser Phe Phe Ala Leu Arg Asp Gln Ile Pro Glu
370 375 380
Leu Glu Asn Asn Glu Lys Ala Pro Lys Val Val Ile Leu Lys Lys Ala
385 390 395 400
Thr Ala Tyr Ile Leu Ser Val Gln Ala Glu Glu Gln Lys Leu Ile Ser
405 410 415
Glu Glu Asp Leu Leu Arg Lys Arg Arg Glu Gln Leu Lys His Lys Leu
420 425 430
Glu Gln Leu Arg Asn Ser Cys Ala
435 440
<210> 11
<211> 620
<212> PRT
<213> Homo sapiens
<400> 11
Met Ala Glu Ala Arg Thr Ser Leu Ser Ala His Cys Arg Gly Pro Leu
1 5 10 15
Ala Thr Gly Leu His Pro Asp Leu Asp Leu Pro Gly Arg Ser Leu Ala
20 25 30
Thr Pro Ala Pro Ser Cys Tyr Leu Leu Gly Ser Glu Pro Ser Ser Gly
35 40 45
Leu Gly Leu Gln Pro Glu Thr His Leu Pro Glu Gly Ser Leu Lys Arg
50 55 60
Cys Cys Val Leu Gly Leu Pro Pro Thr Ser Pro Ala Ser Ser Ser Pro
65 70 75 80
Cys Ala Ser Ser Asp Val Thr Ser Ile Ile Arg Ser Ser Gln Thr Ser
85 90 95
Leu Val Thr Cys Val Asn Gly Leu Arg Ser Pro Pro Leu Thr Gly Asp
100 105 110
Leu Gly Gly Pro Ser Lys Arg Ala Arg Pro Gly Pro Ala Ser Thr Asp
115 120 125
Ser His Glu Gly Ser Leu Gln Leu Glu Ala Cys Arg Lys Ala Ser Phe
130 135 140
Leu Lys Gln Glu Pro Ala Asp Glu Phe Ser Glu Leu Phe Gly Pro His
145 150 155 160
Gln Gln Gly Leu Pro Pro Pro Tyr Pro Leu Ser Gln Leu Pro Pro Gly
165 170 175
Pro Ser Leu Gly Gly Leu Gly Leu Gly Leu Ala Gly Arg Val Val Ala
180 185 190
Gly Arg Gln Ala Cys Arg Trp Val Asp Cys Cys Ala Ala Tyr Glu Gln
195 200 205
Gln Glu Glu Leu Val Arg His Ile Glu Lys Ser His Ile Asp Gln Arg
210 215 220
Lys Gly Glu Asp Phe Thr Cys Phe Trp Ala Gly Cys Val Arg Arg Tyr
225 230 235 240
Lys Pro Phe Asn Ala Arg Tyr Lys Leu Leu Ile His Met Arg Val His
245 250 255
Ser Gly Glu Lys Pro Asn Lys Cys Met Phe Glu Gly Cys Ser Lys Ala
260 265 270
Phe Ser Arg Leu Glu Asn Leu Lys Ile His Leu Arg Ser His Thr Gly
275 280 285
Glu Lys Pro Tyr Leu Cys Gln His Pro Gly Cys Gln Lys Ala Phe Ser
290 295 300
Asn Ser Ser Asp Arg Ala Lys His Gln Arg Thr His Leu Asp Thr Lys
305 310 315 320
Pro Tyr Ala Cys Gln Ile Pro Gly Cys Ser Lys Arg Tyr Thr Asp Pro
325 330 335
Ser Ser Leu Arg Lys His Val Lys Ala His Ser Ala Lys Glu Gln Gln
340 345 350
Val Arg Lys Lys Leu His Ala Gly Pro Asp Thr Glu Ala Asp Val Leu
355 360 365
Thr Glu Cys Leu Val Leu Gln Gln Leu His Thr Ser Thr Gln Leu Ala
370 375 380
Ala Ser Asp Gly Lys Gly Gly Cys Gly Leu Gly Gln Glu Leu Leu Pro
385 390 395 400
Gly Val Tyr Pro Gly Ser Ile Thr Pro His Asn Gly Leu Ala Ser Gly
405 410 415
Leu Leu Pro Pro Ala His Asp Val Pro Ser Arg His His Pro Leu Asp
420 425 430
Ala Thr Thr Ser Ser His His His Leu Ser Pro Leu Pro Met Ala Glu
435 440 445
Ser Thr Arg Asp Gly Leu Gly Pro Gly Leu Leu Ser Pro Ile Val Ser
450 455 460
Pro Leu Lys Gly Leu Gly Pro Pro Pro Leu Pro Pro Ser Ser Gln Ser
465 470 475 480
His Ser Pro Gly Gly Gln Pro Phe Pro Thr Leu Pro Ser Lys Pro Ser
485 490 495
Tyr Pro Pro Phe Gln Ser Pro Pro Pro Pro Pro Leu Pro Ser Pro Gln
500 505 510
Gly Tyr Gln Gly Ser Phe His Ser Ile Gln Ser Cys Phe Pro Tyr Gly
515 520 525
Asp Cys Tyr Arg Met Ala Glu Pro Ala Ala Gly Gly Asp Gly Leu Val
530 535 540
Gly Glu Thr His Gly Phe Asn Pro Leu Arg Pro Asn Gly Tyr His Ser
545 550 555 560
Leu Ser Thr Pro Leu Pro Ala Thr Gly Tyr Glu Ala Leu Ala Glu Ala
565 570 575
Ser Cys Pro Thr Ala Leu Pro Gln Gln Pro Ser Glu Asp Val Val Ser
580 585 590
Ser Gly Pro Glu Asp Cys Gly Phe Phe Pro Asn Gly Ala Phe Asp His
595 600 605
Cys Leu Gly His Ile Pro Ser Ile Tyr Thr Asp Thr
610 615 620
<210> 12
<211> 305
<212> PRT
<213> Homo sapiens
<400> 12
Met Ser Val Asp Pro Ala Cys Pro Gln Ser Leu Pro Cys Phe Glu Ala
1 5 10 15
Ser Asp Cys Lys Glu Ser Ser Pro Met Pro Val Ile Cys Gly Pro Glu
20 25 30
Glu Asn Tyr Pro Ser Leu Gln Met Ser Ser Ala Glu Met Pro His Thr
35 40 45
Glu Thr Val Ser Pro Leu Pro Ser Ser Met Asp Leu Leu Ile Gln Asp
50 55 60
Ser Pro Asp Ser Ser Thr Ser Pro Lys Gly Lys Gln Pro Thr Ser Ala
65 70 75 80
Glu Lys Ser Val Ala Lys Lys Glu Asp Lys Val Pro Val Lys Lys Gln
85 90 95
Lys Thr Arg Thr Val Phe Ser Ser Thr Gln Leu Cys Val Leu Asn Asp
100 105 110
Arg Phe Gln Arg Gln Lys Tyr Leu Ser Leu Gln Gln Met Gln Glu Leu
115 120 125
Ser Asn Ile Leu Asn Leu Ser Tyr Lys Gln Val Lys Thr Trp Phe Gln
130 135 140
Asn Gln Arg Met Lys Ser Lys Arg Trp Gln Lys Asn Asn Trp Pro Lys
145 150 155 160
Asn Ser Asn Gly Val Thr Gln Lys Ala Ser Ala Pro Thr Tyr Pro Ser
165 170 175
Leu Tyr Ser Ser Tyr His Gln Gly Cys Leu Val Asn Pro Thr Gly Asn
180 185 190
Leu Pro Met Trp Ser Asn Gln Thr Trp Asn Asn Ser Thr Trp Ser Asn
195 200 205
Gln Thr Gln Asn Ile Gln Ser Trp Ser Asn His Ser Trp Asn Thr Gln
210 215 220
Thr Trp Cys Thr Gln Ser Trp Asn Asn Gln Ala Trp Asn Ser Pro Phe
225 230 235 240
Tyr Asn Cys Gly Glu Glu Ser Leu Gln Ser Cys Met Gln Phe Gln Pro
245 250 255
Asn Ser Pro Ala Ser Asp Leu Glu Ala Ala Leu Glu Ala Ala Gly Glu
260 265 270
Gly Leu Asn Val Ile Gln Gln Thr Thr Arg Tyr Phe Ser Thr Pro Gln
275 280 285
Thr Met Asp Leu Phe Leu Asn Tyr Ser Met Asn Met Gln Pro Glu Asp
290 295 300
Val
305
<210> 13
<211> 258
<212> PRT
<213> Homo sapiens
<400> 13
Met Arg Ser Phe Asn Gln Val Ser Ser Ala Pro Gly Gly Ala Ser Lys
1 5 10 15
Gly Gly Gly Glu Glu Pro Gly Lys Leu Pro Glu Pro Ala Glu Glu Glu
20 25 30
Ser Gln Val Leu Arg Gly Thr Gly His Cys Lys Trp Phe Asn Val Arg
35 40 45
Met Gly Phe Gly Phe Ile Ser Met Ile Asn Arg Glu Gly Ser Pro Leu
50 55 60
Asp Ile Pro Val Asp Val Phe Val His Gln Ser Lys Leu Phe Met Glu
65 70 75 80
Gly Phe Arg Ser Leu Lys Glu Gly Glu Pro Val Glu Phe Thr Phe Lys
85 90 95
Lys Ser Ser Lys Gly Leu Glu Ser Ile Arg Val Thr Gly Pro Gly Gly
100 105 110
Ser Pro Cys Leu Gly Ser Glu Arg Arg Pro Lys Gly Lys Thr Leu Gln
115 120 125
Lys Arg Lys Pro Lys Gly Asp Arg Cys Tyr Asn Cys Gly Gly Leu Asp
130 135 140
His His Ala Lys Glu Cys Ser Leu Pro Pro Gln Pro Lys Lys Cys His
145 150 155 160
Tyr Cys Gln Ser Ile Met His Met Val Ala Asn Cys Pro His Lys Asn
165 170 175
Val Ala Gln Pro Pro Ala Ser Ser Gln Gly Arg Gln Glu Ala Glu Ser
180 185 190
Gln Pro Cys Thr Ser Thr Leu Pro Arg Glu Val Gly Gly Gly His Gly
195 200 205
Cys Thr Ser Pro Pro Phe Pro Gln Glu Ala Arg Ala Glu Ile Ser Glu
210 215 220
Arg Ser Gly Arg Ser Pro Gln Glu Ala Ser Ser Thr Lys Ser Ser Ile
225 230 235 240
Ala Pro Glu Glu Gln Ser Lys Lys Gly Pro Ser Val Gln Lys Arg Lys
245 250 255
Lys Thr
<210> 14
<211> 7675
<212> DNA
<213> Venezuelan equine encephalitis virus
<400> 14
gaaagttcac gttgacatcg aggaagacag cccattcctc agagctttgc agcggagctt 60
cccgcagttt gaggtagaag ccaagcaggt cactgataat gaccatgcta atgccagagc 120
gttttcgcat ctggcttcaa aactgatcga aacggaggtg gacccatccg acacgatcct 180
tgacattgga agtgcgcccg cccgcagaat gtattctaag cacaagtatc attgtatctg 240
tccgatgaga tgtgcggaag atccggacag attgtataag tatgcaacta agctgaagaa 300
aaactgtaag gaaataactg ataaggaatt ggacaagaaa atgaaggagc tcgccgccgt 360
catgagcgac cctgacctgg aaactgagac tatgtgcctc cacgacgacg agtcgtgtcg 420
ctacgaaggg caagtcgctg tttaccagga tgtatacgcg gttgacggac cgacaagtct 480
ctatcaccaa gccaataagg gagttagagt cgcctactgg ataggctttg acaccacccc 540
ttttatgttt aagaacttgg ctggagcata tccatcatac tctaccaact gggccgacga 600
aaccgtgtta acggctcgta acataggcct atgcagctct gacgttatgg agcggtcacg 660
tagagggatg tccattctta gaaagaagta tttgaaacca tccaacaatg ttctattctc 720
tgttggctcg accatctacc acgagaagag ggacttactg aggagctggc acctgccgtc 780
tgtatttcac ttacgtggca agcaaaatta cacatgtcgg tgtgagacta tagttagttg 840
cgacgggtac gtcgttaaaa gaatagctat cagtccaggc ctgtatggga agccttcagg 900
ctatgctgct acgatgcacc gcgagggatt cttgtgctgc aaagtgacag acacattgaa 960
cggggagagg gtctcttttc ccgtgtgcac gtatgtgcca gctacattgt gtgaccaaat 1020
gactggcata ctggcaacag atgtcagtgc ggacgacgcg caaaaactgc tggttgggct 1080
caaccagcgt atagtcgtca acggtcgcac ccagagaaac accaatacca tgaaaaatta 1140
ccttttgccc gtagtggccc aggcatttgc taggtgggca aaggaatata aggaagatca 1200
agaagatgaa aggccactag gactacgaga tagacagtta gtcatggggt gttgttgggc 1260
ttttagaagg cacaagataa catctattta taagcgcccg gatacccaaa ccatcatcaa 1320
agtgaacagc gatttccact cattcgtgct gcccaggata ggcagtaaca cattggagat 1380
cgggctgaga acaagaatca ggaaaatgtt agaggagcac aaggagccgt cacctctcat 1440
taccgccgag gacgtacaag aagctaagtg cgcagccgat gaggctaagg aggtgcgtga 1500
agccgaggag ttgcgcgcag ctctaccacc tttggcagct gatgttgagg agcccactct 1560
ggaagccgat gtcgacttga tgttacaaga ggctggggcc ggctcagtgg agacacctcg 1620
tggcttgata aaggttacca gctacgctgg cgaggacaag atcggctctt acgctgtgct 1680
ttctccgcag gctgtactca agagtgaaaa attatcttgc atccaccctc tcgctgaaca 1740
agtcatagtg ataacacact ctggccgaaa agggcgttat gccgtggaac cataccatgg 1800
taaagtagtg gtgccagagg gacatgcaat acccgtccag gactttcaag ctctgagtga 1860
aagtgccacc attgtgtaca acgaacgtga gttcgtaaac aggtacctgc accatattgc 1920
cacacatgga ggagcgctga acactgatga agaatattac aaaactgtca agcccagcga 1980
gcacgacggc gaatacctgt acgacatcga caggaaacag tgcgtcaaga aagaactagt 2040
cactgggcta gggctcacag gcgagctggt ggatcctccc ttccatgaat tcgcctacga 2100
gagtctgaga acacgaccag ccgctcctta ccaagtacca accatagggg tgtatggcgt 2160
gccaggatca ggcaagtctg gcatcattaa aagcgcagtc accaaaaaag atctagtggt 2220
gagcgccaag aaagaaaact gtgcagaaat tataagggac gtcaagaaaa tgaaagggct 2280
ggacgtcaat gccagaactg tggactcagt gctcttgaat ggatgcaaac accccgtaga 2340
gaccctgtat attgacgaag cttttgcttg tcatgcaggt actctcagag cgctcatagc 2400
cattataaga cctaaaaagg cagtgctctg cggggatccc aaacagtgcg gtttttttaa 2460
catgatgtgc ctgaaagtgc attttaacca cgagatttgc acacaagtct tccacaaaag 2520
catctctcgc cgttgcacta aatctgtgac ttcggtcgtc tcaaccttgt tttacgacaa 2580
aaaaatgaga acgacgaatc cgaaagagac taagattgtg attgacacta ccggcagtac 2640
caaacctaag caggacgatc tcattctcac ttgtttcaga gggtgggtga agcagttgca 2700
aatagattac aaaggcaacg aaataatgac ggcagctgcc tctcaagggc tgacccgtaa 2760
aggtgtgtat gccgttcggt acaaggtgaa tgaaaatcct ctgtacgcac ccacctcaga 2820
acatgtgaac gtcctactga cccgcacgga ggaccgcatc gtgtggaaaa cactagccgg 2880
cgacccatgg ataaaaacac tgactgccaa gtaccctggg aatttcactg ccacgataga 2940
ggagtggcaa gcagagcatg atgccatcat gaggcacatc ttggagagac cggaccctac 3000
cgacgtcttc cagaataagg caaacgtgtg ttgggccaag gctttagtgc cggtgctgaa 3060
gaccgctggc atagacatga ccactgaaca atggaacact gtggattatt ttgaaacgga 3120
caaagctcac tcagcagaga tagtattgaa ccaactatgc gtgaggttct ttggactcga 3180
tctggactcc ggtctatttt ctgcacccac tgttccgtta tccattagga ataatcactg 3240
ggataactcc ccgtcgccta acatgtacgg gctgaataaa gaagtggtcc gtcagctctc 3300
tcgcaggtac ccacaactgc ctcgggcagt tgccactgga agagtctatg acatgaacac 3360
tggtacactg cgcaattatg atccgcgcat aaacctagta cctgtaaaca gaagactgcc 3420
tcatgcttta gtcctccacc ataatgaaca cccacagagt gacttttctt cattcgtcag 3480
caaattgaag ggcagaactg tcctggtggt cggggaaaag ttgtccgtcc caggcaaaat 3540
ggttgactgg ttgtcagacc ggcctgaggc taccttcaga gctcggctgg atttaggcat 3600
cccaggtgat gtgcccaaat atgacataat atttgttaat gtgaggaccc catataaata 3660
ccatcactat cagcagtgtg aagaccatgc cattaagctt agcatgttga ccaagaaagc 3720
ttgtctgcat ctgaatcccg gcggaacctg tgtcagcata ggttatggtt acgctgacag 3780
ggccagcgaa agcatcattg gtgctatagc gcggcagttc aagttttccc gggtatgcaa 3840
accgaaatcc tcacttgaag agacggaagt tctgtttgta ttcattgggt acgatcgcaa 3900
ggcccgtacg cacaatcctt acaagctttc atcaaccttg accaacattt atacaggttc 3960
cagactccac gaagccggat gtgcaccctc atatcatgtg gtgcgagggg atattgccac 4020
ggccaccgaa ggagtgatta taaatgctgc taacagcaaa ggacaacctg gcggaggggt 4080
gtgcggagcg ctgtataaga aattcccgga aagcttcgat ttacagccga tcgaagtagg 4140
aaaagcgcga ctggtcaaag gtgcagctaa acatatcatt catgccgtag gaccaaactt 4200
caacaaagtt tcggaggttg aaggtgacaa acagttggca gaggcttatg agtccatcgc 4260
taagattgtc aacgataaca attacaagtc agtagcgatt ccactgttgt ccaccggcat 4320
cttttccggg aacaaagatc gactaaccca atcattgaac catttgctga cagctttaga 4380
caccactgat gcagatgtag ccatatactg cagggacaag aaatgggaaa tgactctcaa 4440
ggaagcagtg gctaggagag aagcagtgga ggagatatgc atatccgacg actcttcagt 4500
gacagaacct gatgcagagc tggtgagggt gcatccgaag agttctttgg ctggaaggaa 4560
gggctacagc acaagcgatg gcaaaacttt ctcatatttg gaagggacca agtttcacca 4620
ggcggccaag gatatagcag aaattaatgc catgtggccc gttgcaacgg aggccaatga 4680
gcaggtatgc atgtatatcc tcggagaaag catgagcagt attaggtcga aatgccccgt 4740
cgaagagtcg gaagcctcca caccacctag cacgctgcct tgcttgtgca tccatgccat 4800
gactccagaa agagtacagc gcctaaaagc ctcacgtcca gaacaaatta ctgtgtgctc 4860
atcctttcca ttgccgaagt atagaatcac tggtgtgcag aagatccaat gctcccagcc 4920
tatattgttc tcaccgaaag tgcctgcgta tattcatcca aggaagtatc tcgtggaaac 4980
accaccggta gacgagactc cggagccatc ggcagagaac caatccacag aggggacacc 5040
tgaacaacca ccacttataa ccgaggatga gaccaggact agaacgcctg agccgatcat 5100
catcgaagag gaagaagagg atagcataag tttgctgtca gatggcccga cccaccaggt 5160
gctgcaagtc gaggcagaca ttcacgggcc gccctctgta tctagctcat cctggtccat 5220
tcctcatgca tccgactttg atgtggacag tttatccata cttgacaccc tggagggagc 5280
tagcgtgacc agcggggcaa cgtcagccga gactaactct tacttcgcaa agagtatgga 5340
gtttctggcg cgaccggtgc ctgcgcctcg aacagtattc aggaaccctc cacatcccgc 5400
tccgcgcaca agaacaccgt cacttgcacc cagcagggcc tgctcgagaa ccagcctagt 5460
ttccaccccg ccaggcgtga atagggtgat cactagagag gagctcgagg cgcttacccc 5520
gtcacgcact cctagcaggt cggtctcgag aaccagcctg gtctccaacc cgccaggcgt 5580
aaatagggtg attacaagag aggagtttga ggcgttcgta gcacaacaac aatgacggtt 5640
tgatgcgggt gcatacatct tttcctccga caccggtcaa gggcatttac aacaaaaatc 5700
agtaaggcaa acggtgctat ccgaagtggt gttggagagg accgaattgg agatttcgta 5760
tgccccgcgc ctcgaccaag aaaaagaaga attactacgc aagaaattac agttaaatcc 5820
cacacctgct aacagaagca gataccagtc caggaaggtg gagaacatga aagccataac 5880
agctagacgt attctgcaag gcctagggca ttatttgaag gcagaaggaa aagtggagtg 5940
ctaccgaacc ctgcatcctg ttcctttgta ttcatctagt gtgaaccgtg ccttttcaag 6000
ccccaaggtc gcagtggaag cctgtaacgc catgttgaaa gagaactttc cgactgtggc 6060
ttcttactgt attattccag agtacgatgc ctatttggac atggttgacg gagcttcatg 6120
ctgcttagac actgccagtt tttgccctgc aaagctgcgc agctttccaa agaaacactc 6180
ctatttggaa cccacaatac gatcggcagt gccttcagcg atccagaaca cgctccagaa 6240
cgtcctggca gctgccacaa aaagaaattg caatgtcacg caaatgagag aattgcccgt 6300
attggattcg gcggccttta atgtggaatg cttcaagaaa tatgcgtgta ataatgaata 6360
ttgggaaacg tttaaagaaa accccatcag gcttactgaa gaaaacgtgg taaattacat 6420
taccaaatta aaaggaccaa aagctgctgc tctttttgcg aagacacata atttgaatat 6480
gttgcaggac ataccaatgg acaggtttgt aatggactta aagagagacg tgaaagtgac 6540
tccaggaaca aaacatactg aagaacggcc caaggtacag gtgatccagg ctgccgatcc 6600
gctagcaaca gcgtatctgt gcggaatcca ccgagagctg gttaggagat taaatgcggt 6660
cctgcttccg aacattcata cactgtttga tatgtcggct gaagactttg acgctattat 6720
agccgagcac ttccagcctg gggattgtgt tctggaaact gacatcgcgt cgtttgataa 6780
aagtgaggac gacgccatgg ctctgaccgc gttaatgatt ctggaagact taggtgtgga 6840
cgcagagctg ttgacgctga ttgaggcggc tttcggcgaa atttcatcaa tacatttgcc 6900
cactaaaact aaatttaaat tcggagccat gatgaaatct ggaatgttcc tcacactgtt 6960
tgtgaacaca gtcattaaca ttgtaatcgc aagcagagtg ttgagagaac ggctaaccgg 7020
atcaccatgt gcagcattca ttggagatga caatatcgtg aaaggagtca aatcggacaa 7080
attaatggca gacaggtgcg ccacctggtt gaatatggaa gtcaagatta tagatgctgt 7140
ggtgggcgag aaagcgcctt atttctgtgg agggtttatt ttgtgtgact ccgtgaccgg 7200
cacagcgtgc cgtgtggcag accccctaaa aaggctgttt aagcttggca aacctctggc 7260
agcagacgat gaacatgatg atgacaggag aagggcattg catgaagagt caacacgctg 7320
gaaccgagtg ggtattcttt cagagctgtg caaggcagta gaatcaaggt atgaaaccgt 7380
aggaacttcc atcatagtta tggccatgac tactctagct agcagtgtta aatcattcag 7440
ctacctgaga ggggccccta taactctcta cggctaacct gaatggacta cgacatagtc 7500
tagtccgcca agtctagcat atgggcgcgt gaattcgccg cgaattggca agctgcttac 7560
atagaactcg cggcgattgg catgccgcct taaaattttt attttatttt tcttttcttt 7620
tccgaatcgg attttgtttt taatatttca aaaaaaaaaa aaaaaaaaaa aaaaa 7675
<210> 15
<211> 7675
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<223> /note="Description of Artificial Sequence: Synthetic
polynucleotide"
<400> 15
gaaagttcac gttgacatcg aggaagacag cccattcctc agagctttgc agcggagctt 60
cccgcagttt gaggtagaag ccaagcaggt cactgataat gaccatgcta atgccagagc 120
gttttcgcat ctggcttcaa aactgatcga aacggaggtg gacccatccg acacgatcct 180
tgacattgga agtgcgcccg cccgcagaat gtattctaag cacaagtatc attgtatctg 240
tccgatgaga tgtgcggaag atccggacag attgtataag tatgcaacta agctgaagaa 300
aaactgtaag gaaataactg ataaggaatt ggacaagaaa atgaaggagc tggccgccgt 360
catgagcgac cctgacctgg aaactgagac tatgtgcctc cacgacgacg agtcgtgtcg 420
ctacgaaggg caagtcgctg tttaccagga tgtatacgcg gttgacggac cgacaagtct 480
ctatcaccaa gccaataagg gagttagagt cgcctactgg ataggctttg acaccacccc 540
ttttatgttt aagaacttgg ctggagcata tccatcatac tctaccaact gggccgacga 600
aaccgtgtta acggctcgta acataggcct atgcagctct gacgttatgg agcggtcacg 660
tagagggatg tccattctta gaaagaagta tttgaaacca tccaacaatg ttctattctc 720
tgttggctcg accatctacc acgagaagag ggacttactg aggagctggc acctgccgtc 780
tgtatttcac ttacgtggca agcaaaatta cacatgtcgg tgtgagacta tagttagttg 840
cgacgggtac gtcgttaaaa gaatagctat cagtccaggc ctgtatggga agccttcagg 900
ctatgctgct acgatgcacc gcgagggatt cttgtgctgc aaagtgacag acacattgaa 960
cggggagagg gtctcttttc ccgtgtgcac gtatgtgcca gctacattgt gtgaccaaat 1020
gactggcata ctggcaacag atgtcagtgc ggacgacgcg caaaaactgc tggttgggct 1080
caaccagcgt atagtcgtca acggtcgcac ccagagaaac accaatacca tgaaaaatta 1140
ccttttgccc gtagtggccc aggcatttgc taggtgggca aaggaatata aggaagatca 1200
agaagatgaa aggccactag gactacgaga tagacagtta gtcatggggt gttgttgggc 1260
ttttagaagg cacaagataa catctattta taagcgcccg gatacccaaa ccatcatcaa 1320
agtgaacagc gatttccact cattcgtgct gcccaggata ggcagtaaca cattggagat 1380
cgggctgaga acaagaatca ggaaaatgtt agaggagcac aaggagccgt cacctctcat 1440
taccgccgag gacgtacaag aagctaagtg cgcagccgat gaggctaagg aggtgcgtga 1500
agccgaggag ttgcgcgcag ctctaccacc tttggcagct gatgttgagg agcccactct 1560
ggaggcagac gtcgacttga tgttacaaga ggctggggcc ggctcagtgg agacacctcg 1620
tggcttgata aaggttacca gctacgatgg cgaggacaag atcggctctt acgctgtgct 1680
ttctccgcag gctgtactca agagtgaaaa attatcttgc atccaccctc tcgctgaaca 1740
agtcatagtg ataacacact ctggccgaaa agggcgttat gccgtggaac cataccatgg 1800
taaagtagtg gtgccagagg gacatgcaat acccgtccag gactttcaag ctctgagtga 1860
aagtgccacc attgtgtaca acgaacgtga gttcgtaaac aggtacctgc accatattgc 1920
cacacatgga ggagcgctga acactgatga agaatattac aaaactgtca agcccagcga 1980
gcacgacggc gaatacctgt acgacatcga caggaaacag tgcgtcaaga aagaactagt 2040
cactgggcta gggctcacag gcgagctggt ggatcctccc ttccatgaat tcgcctacga 2100
gagtctgaga acacgaccag ccgctcctta ccaagtacca accatagggg tgtatggcgt 2160
gccaggatca ggcaagtctg gcatcattaa aagcgcagtc accaaaaaag atctagtggt 2220
gagcgccaag aaagaaaact gtgcagaaat tataagggac gtcaagaaaa tgaaagggct 2280
ggacgtcaat gccagaactg tggactcagt gctcttgaat ggatgcaaac accccgtaga 2340
gaccctgtat attgacgaag cttttgcttg tcatgcaggt actctcagag cgctcatagc 2400
cattataaga cctaaaaagg cagtgctctg cggggatccc aaacagtgcg gtttttttaa 2460
catgatgtgc ctgaaagtgc attttaacca cgagatttgc acacaagtct tccacaaaag 2520
catctctcgc cgttgcacta aatctgtgac ttcggtcgtc tcaaccttgt tttacgacaa 2580
aaaaatgaga acgacgaatc cgaaagagac taagattgtg attgacacta ccggcagtac 2640
caaacctaag caggacgatc tcattctcac ttgtttcaga gggtgggtga agcagttgca 2700
aatagattac aaaggcaacg aaataatgac ggcagctgcc tctcaagggc tgacccgtaa 2760
aggtgtgtat gccgttcggt acaaggtgaa tgaaaatcct ctgtacgcac ccacctcaga 2820
acatgtgaac gtcctactga cccgcacgga ggaccgcatc gtgtggaaaa cactagccgg 2880
cgacccatgg ataaaaacac tgactgccaa gtaccctggg aatttcactg ccacgataga 2940
ggagtggcaa gcagagcatg atgccatcat gaggcacatc ttggagagac cggaccctac 3000
cgacgtcttc cagaataagg caaacgtgtg ttgggccaag gctttagtgc cggtgctgaa 3060
gaccgctggc atagacatga ccactgaaca atggaacact gtggattatt ttgaaacgga 3120
caaagctcac tcagcagaga tagtattgaa ccaactatgc gtgaggttct ttggactcga 3180
tctggactcc ggtctatttt ctgcacccac tgttccgtta tccattagga ataatcactg 3240
ggataactcc ccgtcgccta acatgtacgg gctgaataaa gaagtggtcc gtcagctctc 3300
tcgcaggtac ccacaactgc ctcgggcagt tgccactgga agagtctatg acatgaacac 3360
tggtacactg cgcaattatg atccgcgcat aaacctagta cctgtaaaca gaagactgcc 3420
tcatgcttta gtcctccacc ataatgaaca cccacagagt gacttttctt cattcgtcag 3480
caaattgaag ggcagaactg tcctggtggt cggggaaaag ttgtccgtcc caggcaaaat 3540
ggttgactgg ttgtcagacc ggcctgaggc taccttcaga gctcggctgg atttaggcat 3600
cccaggtgat gtgcccaaat atgacataat atttgttaat gtgaggaccc catataaata 3660
ccatcactat cagcagtgtg aagaccatgc cattaagctt agcatgttga ccaagaaagc 3720
ttgtctgcat ctgaatcccg gcggaacctg tgtcagcata ggttatggtt acgctgacag 3780
ggccagcgaa agcatcattg gtgctatagc gcggcagttc aagttttccc gggtatgcaa 3840
accgaaatcc tcacttgaag agacggaagt tctgtttgta ttcattgggt acgatcgcaa 3900
ggcccgtacg cacaattctt acaagctttc atcaaccttg accaacattt atacaggttc 3960
cagactccac gaagccggat gtgcaccctc atatcatgtg gtgcgagggg atattgccac 4020
ggccaccgaa ggagtgatta taaatgctgc taacagcaaa ggacaacctg gcggaggggt 4080
gtgcggagcg ctgtataaga aattcccgga aagcttcgat ttacagccga tcgaagtagg 4140
aaaagcgcga ctggtcaaag gtgcagctaa acatatcatt catgccgtag gaccaaactt 4200
caacaaagtt tcggaggttg aaggtgacaa acagttggca gaggcttatg agtccatcgc 4260
taagattgtc aacgataaca attacaagtc agtagcgatt ccactgttgt ccaccggcat 4320
cttttccggg aacaaagatc gactaaccca atcattgaac catttgctga cagctttaga 4380
caccactgat gcagatgtag ccatatactg cagggacaag aaatgggaaa tgactctcaa 4440
ggaagcagtg gctaggagag aagcagtgga ggagatatgc atatccgacg actcttcagt 4500
gacagaacct gatgcagagc tggtgagggt gcatccgaag agttctttgg ctggaaggaa 4560
gggctacagc acaagcgatg gcaaaacttt ctcatatttg gaagggacca agtttcacca 4620
ggcggccaag gatatagcag aaattaatgc catgtggccc gttgcaacgg aggccaatga 4680
gcaggtatgc atgtatatcc tcggagaaag catgagcagt attaggtcga aatgccccgt 4740
cgaagagtcg gaagcctcca caccacctag cacgctgcct tgcttgtgca tccatgccat 4800
gactccagaa agagtacagc gcctaaaagc ctcacgtcca gaacaaatta ctgtgtgctc 4860
atcctttcca ttgccgaagt atagaatcac tggtgtgcag aagatccaat gctcccagcc 4920
tatattgttc tcaccgaaag tgcctgcgta tattcatcca aggaagtatc tcgtggaaac 4980
accaccggta gacgagactc cggagccatc ggcagagaac caatccacag aggggacacc 5040
tgaacaacca ccacttataa ccgaggatga gaccaggact agaacgcctg agccgatcat 5100
catcgaagag gaagaagagg atagcataag tttgctgtca gatggcccga cccaccaggt 5160
gctgcaagtc gaggcagaca ttcacgggcc gccctctgta tctagctcat cctggtccat 5220
tcctcatgca tccgactttg atgtggacag tttatccata cttgacaccc tggagggagc 5280
tagcgtgacc agcggggcaa cgtcagccga gactaactct tacttcgcaa agagtatgga 5340
gtttctggcg cgaccggtgc ctgcgcctcg aacagtattc aggaaccctc cacatcccgc 5400
tccgcgcaca agaacaccgt cacttgcacc cagcagggcc tgctcgagaa ccagcctagt 5460
ttccaccccg ccaggcgtga atagggtgat cactagagag gagctcgagg cgcttacccc 5520
gtcacgcact cctagcaggt cggtctcgag aaccagcctg gtctccaacc cgccaggcgt 5580
aaatagggtg attacaagag aggagtttga ggcgttcgta gcacaacaac aatgacggtt 5640
tgatgcgggt gcatacatct tttcctccga caccggtcaa gggcatttac aacaaaaatc 5700
agtaaggcaa acggtgctat ccgaagtggt gttggagagg accgaattgg agatttcgta 5760
tgccccgcgc ctcgaccaag aaaaagaaga attactacgc aagaaattac agttaaatcc 5820
cacacctgct aacagaagca gataccagtc caggaaggtg gagaacatga aagccataac 5880
agctagacgt attctgcaag gcctagggca ttatttgaag gcagaaggaa aagtggagtg 5940
ctaccgaacc ctgcatcctg ttcctttgta ttcatctagt gtgaaccgtg ccttttcaag 6000
ccccaaggtc gcagtggaag cctgtaacgc catgttgaaa gagaactttc cgactgtggc 6060
ttcttactgt attattccag agtacgatgc ctatttggac atggttgacg gagcttcatg 6120
ctgcttagac actgccagtt tttgccctgc aaagctgcgc agctttccaa agaaacactc 6180
ctatttggaa cccacaatac gatcggcagt gccttcagcg atccagaaca cgctccagaa 6240
cgtcctggca gctgccacaa aaagaaattg caatgtcacg caaatgagag aattgcccgt 6300
attggattcg gcggccttta atgtggaatg cttcaagaaa tatgcgtgta ataatgaata 6360
ttgggaaacg tttaaagaaa accccatcag gcttactgaa gaaaacgtgg taaattacat 6420
taccaaatta aaaggaccaa aagctgctgc tctttttgcg aagacacata atttgaatat 6480
gttgcaggac ataccaatgg acaggtttgt aatggactta aagagagacg tgaaagtgac 6540
tccaggaaca aaacatactg aagaacggcc caaggtacag gtgatccagg ctgccgatcc 6600
gctagcaaca gcgtatctgt gcggaatcca ccgagagctg gttaggagat taaatgcggt 6660
cctgcttccg aacattcata cactgtttga tatgtcggct gaagactttg acgctattat 6720
agccgagcac ttccagcctg gggattgtgt tctggaaact gacatcgcgt cgtttgataa 6780
aagtgaggac gacgccatgg ctctgaccgc gttaatgatt ctggaagact taggtgtgga 6840
cgcagagctg ttgacgctga ttgaggcggc tttcggcgaa atttcatcaa tacatttgcc 6900
cactaaaact aaatttaaat tcggagccat gatgaaatct ggaatgttcc tcacactgtt 6960
tgtgaacaca gtcattaaca ttgtaatcgc aagcagagtg ttgagagaac ggctaaccgg 7020
atcaccatgt gcagcattca ttggagatga caatatcgtg aaaggagtca aatcggacaa 7080
attaatggca gacaggtgcg ccacctggtt gaatatggaa gtcaagatta tagatgctgt 7140
ggtgggcgag aaagcgcctt atttctgtgg agggtttatt ttgtgtgact ccgtgaccgg 7200
cacagcgtgc cgtgtggcag accccctaaa aaggctgttt aagcttggca aacctctggc 7260
agcagacgat gaacatgatg atgacaggag aagggcattg catgaagagt caacacgctg 7320
gaaccgagtg ggtattcttt cagagctgtg caaggcagta gaatcaaggt atgaaaccgt 7380
aggaacttcc atcatagtta tggccatgac tactctagct agcagtgtta aatcattcag 7440
ctacctgaga ggggccccta taactctcta cggctaacct gaatggacta cgacatagtc 7500
tagtccgcca agtctagcat atgggcgcgt gaattcgccg cgaattggca agctgcttac 7560
atagaactcg cggcgattgg catgccgcct taaaattttt attttatttt tcttttcttt 7620
tccgaatcgg attttgtttt taatatttca aaaaaaaaaa aaaaaaaaaa aaaaa 7675
Claims (30)
- 조혈 계통의 출발 세포로부터 유도 만능 줄기 세포 (iPSC) 집단을 수득하는 방법으로서,
BCL-xL, 및 OCT 패밀리 단백질, KLF 패밀리 단백질, MYC 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물을 출발 세포에 도입하는 단계, 및
출발 세포를 배양하여 BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자를 발현시켜 출발 세포 및 그의 자손이 iPSC로 재프로그래밍되도록 유도하는 단계를 포함하는 방법. - 제1항에 있어서, 출발 세포가 인간 기원의 조혈 줄기 세포, 적혈구 전구 세포, 림프양 전구 세포, 말초 혈액 단핵 세포, T 림프구, B 림프구, 대식세포, 단핵구, 호중구, 호산구, 또는 수지상 세포인 방법.
- 제2항에 있어서, 출발 세포가 에리트로포이에틴, 줄기 세포 인자, 및 IL-3의 존재하에 임의적으로 5 내지 10일, 또는 6 내지 7일 동안 말초 혈액 단핵 세포 (PBMC)를 배양함으로써 수득된 적혈구 전구 세포인 방법.
- 제3항에 있어서, PBMC가 0.5-5 IU/ml 에리트로포이에틴, 50-200 ng/mL 줄기 세포 인자, 및 1-10 ng/mL IL-3의 존재하에 배양되는 것인 방법.
- 제1항 내지 제5항 중 어느 한 항에 있어서, RNA 발현 구축물이 전기천공을 통해 출발 세포에 도입되는 것인 방법.
- 제5항에 있어서, 출발 세포를 전기천공 전에 B18R 단백질과 접촉시키는 단계를 포함하는 방법.
- 제1항 내지 제6항 중 어느 한 항의 방법으로부터 수득된 유도 만능 줄기 세포 (iPSC) 집단.
- BCL-xL, 및 Oct 패밀리 단백질, KLF 패밀리 단백질, Myc 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물로 형질감염된 조혈 계통의 출발 세포로부터 수득된 유도 만능 줄기 세포 (iPSC) 집단.
- 제8항에 있어서, 출발 세포가 인간 기원의 조혈 줄기 세포, 적혈구 전구 세포, 림프양 전구 세포, 말초 혈액 단핵 세포, T 림프구, B 림프구, 대식세포, 단핵구, 호중구, 호산구, 또는 수지상 세포인 iPSC.
- 제9항에 있어서, 출발 세포가 EPO, SCF, 및 IL-3의 존재하에 임의적으로 5 내지 10일, 또는 6 내지 7일 동안 말초 혈액 단핵 세포 (PBMC)를 배양함으로써 수득된 적혈구 전구 세포인 iPSC.
- BCL-xL, 및 OCT 패밀리 단백질, KLF 패밀리 단백질, MYC 패밀리 단백질, SOX 패밀리 단백질, LIN28 단백질, NANOG 단백질, 및 p53 우성 음성 단백질로부터 선택되는 하나 이상의 추가의 재프로그래밍 인자를 코딩하는 알파바이러스 RNA 발현 구축물.
- 제11항의 알파바이러스 RNA 발현 구축물에 대한 코딩 서열을 포함하는 DNA 벡터.
- 제11항의 알파바이러스 RNA 발현 구축물 또는 제12항의 DNA 벡터를 포함하는 숙주 세포로서, 임의적으로 여기서 숙주 세포는 인간 세포인 숙주 세포.
- 제1항 내지 제13항 중 어느 한 항에 있어서, 알파바이러스 RNA 발현 구축물이 자기 복제성이고, nsP1, nsP2, nsP3, 및 nsP4 유전자를 포함하는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포.
- 제1항 내지 제14항 중 어느 한 항에 있어서, 알파바이러스 RNA 발현 구축물이 베네수엘라 말 뇌염 바이러스 (VEEV) RNA 발현 구축물이고, VEEV nsP1, nsP2, nsP3, 및 nsP4 유전자를 포함하고,
임의적으로 여기서 VEEV RNA 발현 구축물이 야생형 VEEV 게놈의 상응하는 영역(들)으로부터 하나 이상의, 임의적으로 2개 이상, 3개 이상, 4개 이상, 5개 이상, 또는 6개 이상의 돌연변이를 함유하는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포. - 제1항 내지 제15항 중 어느 한 항에 있어서,
OCT 패밀리 단백질이 OCT4, 임의적으로, 인간 OCT4이고/거나,
KLF 패밀리 단백질이 KLF4, 임의적으로, 인간 KLF4이고/거나,
SOX 패밀리 단백질이 SOX2, 임의적으로, 인간 SOX2이고/거나,
LIN28 단백질이 LIN28B, 임의적으로, 인간 LIN28B이고/거나,
MYC 패밀리 단백질이 c-MYC, 임의적으로, 인간 c-MYC인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포. - 제1항 내지 제16항 중 어느 한 항에 있어서, 알파바이러스 RNA 발현 구축물이
서열식별번호(SEQ ID NO:) 1 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 BCL-xL 단백질,
서열식별번호 3 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 OCT4 단백질,
서열식별번호 5 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 KLF4 단백질,
서열식별번호 7 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 SOX2 단백질, 및/또는
서열식별번호 9 또는 그와 적어도 95% 동일한 아미노산 서열을 포함하는 c-MYC 단백질의 발현을 지시하는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포. - 제1항 내지 제17항 중 어느 한 항에 있어서, 알파바이러스 RNA 발현 구축물이 OCT 패밀리 단백질, SOX 패밀리 단백질, BCL-xL, 및 MYC 패밀리 단백질, 및 임의적으로 KLF 패밀리 단백질의 발현을 지시하는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포.
- 제1항 내지 제18항 중 어느 한 항에 있어서, BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자에 대한 코딩 서열이 2A 펩티드에 대한 코딩 서열 또는 내부 리보솜 진입 부위 (IRES)에 의해 이격되어 있는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포.
- 제1항 내지 제19항 중 어느 한 항에 있어서, BCL-xL 및 하나 이상의 추가의 재프로그래밍 인자에 대한 코딩 서열이 공통 프로모터, 임의적으로, 26S 프로모터의 전사 제어하에 있는 것인 방법, iPSC, 알파바이러스 RNA 발현 구축물, DNA 벡터, 또는 숙주 세포.
- 분화 촉진제의 존재하에 제7항 내지 제10항 및 제14항 내지 제20항 중 어느 한 항의 iPSC를 배양하는 단계를 포함하는, 시험관내에서 분화된 세포를 수득하는 방법.
- 제21항의 방법에 의해 수득된 분화된 세포.
- 제7항 내지 제10항 및 제14항 내지 제20항 중 어느 한 항의 iPSC의 분화로부터 수득된 분화된 세포.
- 제21항 내지 제23항 중 어느 한 항에 있어서, 분화된 세포가
임의적으로 T 세포, 키메라 항원 수용체 (CAR)를 발현하는 T 세포, 억제 T 세포, 골수성 세포, 수지상 세포, 및 면역억제 대식세포로부터 선택되는 인간 면역 세포;
임의적으로 도파민성 뉴런, 미세아교세포, 희소돌기아교세포, 성상세포, 피질 뉴런, 척수 또는 안구운동 뉴런, 장 뉴런, 기원판 유래 세포, 슈반 세포, 및 삼차신경 또는 감각 뉴런으로부터 선택되는 인간 신경계 세포;
임의적으로 심근세포, 내피 세포, 및 결절 세포로부터 선택되는 인간 심혈관계 세포;
임의적으로 간세포, 담관세포, 및 췌장 베타 세포로부터 선택되는 인간 대사계 세포, 또는
임의적으로 망막 색소 상피 세포, 광수용체 추상 세포, 광수용체 간상 세포, 양극 세포, 또는 신경절 세포로부터 선택되는 인간 안구계 세포인 방법, 또는 분화된 세포. - 제21항 내지 제24항 중 어느 한 항에 있어서, 분화된 세포가 외배엽 계통의 것이고, 임의적으로 여기서 분화된 세포가 뉴런인 방법 또는 세포.
- 제21항 내지 제24항 중 어느 한 항에 있어서, 분화된 세포가 중배엽 계통의 것이고, 임의적으로 여기서 분화된 세포가 심근세포인 방법 또는 세포.
- 제22항 내지 제26항 중 어느 한 항의 분화된 세포 및 제약상 허용되는 담체를 포함하는 제약 조성물.
- 치료를 필요로 하는 환자에게 제22항 내지 제26항 중 어느 한 항의 분화된 세포 또는 제27항의 제약 조성물을 투여하는 단계를 포함하는, 치료를 필요로 하는 환자를 치료하는 방법.
- 치료를 필요로 하는 환자를 치료하기 위한 의약의 제조를 위한 제22항 내지 제26항 중 어느 한 항의 분화된 세포의 용도.
- 제22항 내지 제27항 중 어느 한 항에 있어서, 치료를 필요로 하는 환자 치료에서 사용하기 위한 분화된 세포, 또는 제약 조성물.
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