KR20150097480A - 항원-특이적인 t 세포 반응을 유도하는 면역 강화제로서 사용을 위한 융합 단백질 - Google Patents
항원-특이적인 t 세포 반응을 유도하는 면역 강화제로서 사용을 위한 융합 단백질 Download PDFInfo
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- KR20150097480A KR20150097480A KR1020157014330A KR20157014330A KR20150097480A KR 20150097480 A KR20150097480 A KR 20150097480A KR 1020157014330 A KR1020157014330 A KR 1020157014330A KR 20157014330 A KR20157014330 A KR 20157014330A KR 20150097480 A KR20150097480 A KR 20150097480A
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Abstract
본 발명은 강화된 항원-특이적인 T 세포 반응을 위한 면역원 강화제(enhancer)로서의 융합 단백질의 사용에 대한 것이다. 상기 융합 단백질은 하기를 포함한다: (a) 항원-제시 세포(antigen-presenting cell; APC)-결합 도메인 또는 CD91 수용체-결합 도메인; (b) 단백질 형질도입(transduction) 도메인; 및 (c) 병원균(pathogen)의 항원, 이는 융합 단백질의 N-말단에 위치한 APC-결합 도메인 또는 CD91 수용체-결합 도메인, 및 단백질 형질도입 도메인의 C-말단에 위치한 병원균의 항원. 상기 단백질 형질도입 도메인은 하기로 구성된 그룹으로부터 선별된 것이다: (i) T 세포 민감성 신호-형질도입 펩타이드, 링커, 및 전좌(translocation) 펩타이드를 포함하는 융합 폴리펩타이드; (ii) T 세포-민감성 신호-형질도입 펩타이드; 및 (iii) 34 내지 112 아미노산 잔기 길이의 전좌 펩타이드.
Description
본 발명은 일반적으로 융합 단백질(fusion protein), 및 더 자세하게는 T 세포-매개(T cell-mediated)의 면역 반응(immune response)을 강화하기 위한 융합 단백질과 연관된 것이다.
분자 생물학(molecular biology)은 면역원(immunogen)이 모단백질(parent protein) 또는 복합체(complex)의 절편 또는 소단위(subunit)인, 소단위 백신(vaccine)의 생산을 가능하게 하였다. T 세포 민감성 반응(T cell sensitizing responses)을 유도할 수 있고 가염성 시약(infectious agent)의 많은 종(strains)으로부터 서열을 포함하기에 충분히 유연한 안정한 백신의 개발이 필요하다.
한 측면에서, 본 발명은 하기를 포함하는 융합 단백질과 연관된다:
(a) 융합 단백질의 N-말단에 위치하는 항원-제시 세포(antigen-presenting cell; APC)-결합 도메인 또는 CD91 수용체-결합 도메인;
(b) APC-결합 도메인 또는 CD91 수용체-결합 도메인의 C-말단에 위치한 단백질 형질도입 도메인, 상기 단백질 형질도입 도메인은 하기로 구성된 군으로부터 선별된다:
(i) T 세포 민감성 신호-형질도입 펩타이드, 링커, 및 전좌(translocation) 펩타이드를 포함하는 융합 폴리펩타이드, 여기서:
(1) 상기 융합 폴리펩타이드의 N-말단에 위치한 T 세포 민감성 신호-형질도입 펩타이드;
(2) 상기 T 세포 민감성 신호-형질도입 펩타이드 및 전좌 펩타이드를 연결하는 서열번호 15를 포함하는 링커; 및
(3) 상기 전좌 펩타이드는 34 내지 112 아미노산 잔기 길이 및 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열을 포함;
(ii) T 세포-민감성 신호-형질도입 펩타이드; 및
(iii) 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열을 포함하는 34 내지 112 아미노산 잔기 길이의 전좌 펩타이드; 및
(c) 단백질 형질도입 도메인의 C-말단에 위치한 병원균의 항원, 여기서:
상기 T 세포-민감성 신호-형질전환 펩타이드는 28 내지 53 아미노사 잔기 길이이고 Xaa8은 I 또는 L; Xaa10은 V, F 또는 A, Xaa11은 M 또는 L, Xaa17은 L 또는 I인 서열번호 31과 최소 90% 동일한 상기 아미노산 서열을 포함하고; 및
상기 APC-결합 도메인 또는 CD91 수용체-결합 도메인은 만약, 단백질 형질도이 도메인이 전좌 펩타이드(iii)인 경우 수도모나스 엑소톡신 A(Pseudomonas exotoxin A; PE) 결합 도메인의 아미노산 서열이 없다.
본 발명의 한 실시예에서, 상기 APC-결합 도메인 또는 CD91 수용체-결합 도메인은 서열번호 5, 9, 6, 7, 및 8로 구성된 군으로부터 선별된 서열과 최소 90% 동일한 아미노산서열을 포함하는 폴리펩타이드이다. 대안적으로, 상기 APC-결합 도메인은 수용체-연관 단백질-1(receptor-associated protein-1; RAP1) 도메인 III, 알파-2-마크로글로불린 수용체-연관 단백질(alpha-2-macroglobulin receptor-associated protein; A2M), HIV-Tat 및 열 충격 단백질(hear shock proteins; HSPs), 및 수도모나스 엑소톡신 A(Pseudomonas exotoxin A; PE) 결합 도메인 I으로 구성되는 군으로부터 선택된 것이다.
본 발명의 또 다른 실시예에서, 상기 융합 단백질은 수도모나스 엑소톡신 A(Pseudomonas exotoxin A; PE) 결합 도메인 I의 아미노산 서열이 없는 것이다.
본 발명의 또 다른 실시예에서, 상기 융합 단백질은 추가적으로 융합 단백질의 C-말단에 위치한 골지체 유지 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 골지체 유지 서열은 Lys-Asp-Glu-Leu(서열번호 14)의 아미노산 서열을 포함한다. 상기 ER 유지 서열은 서열번호 14, 16 내지 19로 이루어진 군으로부터 선택된 서열을 포함할 수 있다. 대안적으로, 상기 ER 유지 서열은 서열번호 16 내지 19로 이루어진 군으로부터 선택된 서열을 구성할 수 있다.
본 발명의 또 다른 실시예에서, 상기 융합 단백질은 만약 항원이 10개 또는 그 이상의 항원결정기(epitope)를 가질 때 그것의 C-말단에 위치한 골지체 유지 서열이 없다.
본 발명의 또 다른 실시예에서, 상기 단백질 형질도입 도메인은 융합 폴리펩타이드(bi)이다.
본 발명의 또 다른 실시예에서, 상기 단백질 형질도입 도메인은 T 세포-민감성 신호-형질도입 펩타이드(bii)이다.
본 발명의 또 다른 실시예에서, 상기 융합 단백질은 추가적으로 서열번호 15를 구성하는, 단백질 형질도입 도메인 및 항원 사이의 추가적인 링커를 포함한다.
본 발명의 또 다른 실시예에서, 상기 단백질 형질도입 도메인은 전좌 펩타이드(biii)이다.
본 발명의 또 다른 실시예에서, 상기 융합 단백질은 추가적으로 서열번호 15를 구성하는, APC-결합 도메인 또는 CD91 수용체-결합 도메인 및 전좌 펩타이드 사이의 추가적인 링커를 포함한다.
본 발명의 또 다른 실시예에서, 상기 단백질 형질도입 도메인은 서열번호 30의 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 APC-결합 도메인은 서열번호 5, 9, 6, 7, 및 8로 이루어진 군으로부터 선택된 서열과 최소 95% 동일한 아미노산 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 APC-결합 도메인은 서열번호 5, 9, 6, 7, 및 8로 이루어진 군으로부터 선택된 아미노산 서열을 포함하는 폴리펩타이드이다.
본 발명의 또 다른 실시예에서, 상기 T 세포민감성 신호-형질도입 펩타이드는 서열번호 1 또는 2와 최소 90% 동일한 아미노산 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 T 세포 민감성 신호-형질도입 펩타이드는 서열번호 1 및 2로 이루어진 군으로부터 선택된 아미노산 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 전좌 펩타이드는 서열번호 3, 20 및 4로 구성된 군으로부터 선택된 상기 아미노산 서열을 포함한다.
본 발명의 또 다른 실시예에서, 상기 전좌 펩타이드는 34 내지 61 아미노산 잔기 길이를 갖는다.
본 발명의 또 다른 실시예에서, 앞서 언급한 것으로서 융합 단백질의 단백질 형질도입 도메인은 하기의 특징을 갖는다: (i) 상기 T 세포-민감성 신호-형질도입 펩타이드는 서열번호 1 또는 2의 아미노산 서열을 포함하고; 및 (ii) 전좌 펩타이드는 서열번호 3과 최소 95% 동일한 아미노산 서열을 포함한다.
상기 T 세포 민감성 신호-형질도입 펩타이드는 (X)2는 I 또는 L; (X)4는 V, F 또는 A, 및 (X)5는 M 또는 L인 T 세포에 있는 CD28 수용체의 K1(X)2E3(X)4(X)5Y6P7P8P9Y10(서열번호 32)의 아미노산 서열을 인식하고 결합하는 항체를 유도하는 특징을 나타낸다.
또 다른 측면에서, 본 발명은 하기로 구성된 융합 단백질과 연관된다:
(a) 융합 단백질의 N-말단에 위치한 항원-제시 세포(APC)-결합 도메인 또는 CD91 수용체-결합 도메인;
(b) APC-결합 도메인 또는 CD91 수용체-결합 도메인의 C-말단에 위치한 단백질 형질도입 도메인, 상기 단백질 형질도입 도메인은 하기로 구성된 군으로부터 선택된다:
(i) T 세포 민감성 신호-형질도입 펩타이드, 링커, 및 전좌 펩타이드로 구성되는 융합 폴리펩타이드, 여기서:
(1) 상기 T 세포 민감성 신호-형질도입 펩타이드는 융합 폴리펩타이드의 N-말단에 위치한다;
(2) 상기 링커는 T 세포 민감성 신호-형질도입 펩타이드 및 전좌 펩타이드를 연결하는 서열번호 15를 포함한다; 및
(3) 상기 전좌 펩타이드는 34 내지 112 아미노산 잔기 길이이고 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열을 포함한다;
(ii) T 세포-민감성 신호-형질도입 펩타이드; 및
(iii) 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열로 구성되는 34 내지 112 아미노산 잔기 길이의 전좌 펩타이드; 및
(c) 단백질 형질도입 도메인의 C-말단에 위치한 병원균의 항원; 여기서:
상기 T 세포-민감성 신호-형질도입 펩타이드는 28 내지 53 아미노산 잔기의 길이이고 Xaa8은 I 또는 L; Xaa10은 V, F 또는 A, Xaa11은 M 또는 L, Xaa17은 L 또는 I인 서열번호 31과 최소 90% 동일한 상기 아미노산 서열을 포함한다; 및
상기 APC-결합 도메인 또는 CD91 수용체-결합 도메인은 만약 단백질 형질도입 도메인이 전좌 펩타이드(biii)인 경우, 수도모나스 엑소톡신 A((Pseudomonas exotoxin A; PE)의 아미노산 서열이 없다.
상기 항원-제시 세포(APC)는 수지상 세포, 대식세포(macrophages), B-세포 및 단핵 백혈구(monocytes)일 수 있다.
본 발명의 한 실시예에서, 상기 APC의 세포 막은 CD91 수용체를 포함한다.
또 다른 측면에서, 본 발명은 하기를 포함하는 백신 조성물과 연관된다: (a) 앞서 말한 것과 같은 융합 단백질의 치료적으로 효과적인 양; 및 (b) 아주번트.
상기 아주번트는 항원 전달 시료 또는 면역 강화제(immune potentiator) 중 하나이다. 본 발명의 한 실시예에서, 상기 백신 조성물은 항원 전달 시표를 포함하고 면역 강화제는 없다.
추가적인 또 다른 측면에서, 본 발명은 앞서 언급한 것으로서 융합 단백질의 치료적으로 효과적인 양을 포함하는 백센 조성물을 그것을 필요로 하는 개체에게 투여하는 것, 및 이로써 강화된 병원균 항원-특이적인 T 세포 반응을 유도하는 것을 포함하는 강화된 병원균 항원-특이적인 T 세포 반응을 유도하기 위한 방법과 연관된다.
추가적인 또 다른 측면에서, 본 발명은 앞서 언급한 것으로서 융합 단백질의 치료적으로 효과적인 양을 포함하는 백센 조성물을 그것을 필요로 하는 개체에게 투여하는 것, 및 이로써 질병 세포의 세포 막에 존재하는 클래스 I 주요 조직적합성 복합체(class I major histocompatibility complex; MHC-I) 분자를 통해 항원에 제시되는 질병 세포를 사멸하는 것을 포함하는, 질병 세포의 세포 막에 존재하는 클래스 I MHC 분자를 통해 항원에 제시되는 질병 세포를 사멸하기 위한 방법과 연관된다.
본 발명의 한 실시예에서, 상기 질병 세포는 암 세포이다.
또 다른 측면에서, 본 발명은 제 1항의 융합 단백질의 치료적으로 효과적인 양을 포함하는 백신 조성물을 그것을 필요로 하는 개체에게 투여하여, 병원균에 의해 야기되는 감염을 예방, 치료하고 및/또는 감염에 의해 야기되는 증상을 최소화하는 것을 포함한다. 본 발명은 또한 강화되 병원균 항원-특이적인 T 세포 반응을 유도하기 위한, 또는 질병 세포의 세포막에 존재하는 클래스 I MHC 분자를 통해 항원에 제시되는 질병 세포를 사멸하기 위한, 또는 병원균에 의해 야기되는 감염의 예방, 치료 및/또는 감염에 의해 야기되는 증상을 최소화에 사용을 위한, 앞서 언급한 것으로서 융합 단백질 또는 백신 조성물과 연관된다.
병원균에 의해 유발된 감염을 예방, 치료하고, 및/ 또는 감염에 의해 야기되는 증상을 최소화하기 위한 융합 단백질과 연관된다.
상기 병원균은 인간 파필로마바이러스(Human Papillomavirus; HPV), 돼지번식호흡장애증후군바이러스(porcine Reproductive and Respiratory Syndrome Virus; PRRSV), 인간면역-결핍바이러스(Human Immuno-deficient virus; HIV-1), 독감 바이러스(flu virus), 댕기 바이러스(Dangue virus), C형 간염 바이러스(Hepatitis C virus; HCV), B형 간염 바이러스(Hepatitis B virus; HBV), 돼지 서코바이러스(Porcine Circovirus 2; PCV2)로 이루어진 군으로부터 선택되는 최소한 하나이다.
본 발명의 한 실시예에서, 앞서 언급한것으로서 상기 융합 단백질은 그것을 필요로하는 개체에 항원-특이적인 세포독성 T 세포 반응을 유도하는데 사용하기 위한 것이다. 상기 융합 단백질은 그것을 필요로 하는 개체에서, 항원-특이적인 CD4+ T 세포 반응을 유도하거나, 또는 강화된 항원-특이적인 항체 역가(titer) 반응을 유도하기 위한 면역원성 강화제로서 사용할 수 있다.
상기 및 다른 측면은 비록 그것에 변형 및 수정이 발명의 신규한 개념의 사상 및 범위를 벗어나지 않고 영향을 받음에도 불구하고, 하기의 도면과 함꼐 바람직한 실시예의 설명으로부터 명백해 질 것이다.
하기 동반되는 도면은 본 발명의 하나 또는 그 이상의 실시예를 설명하고, 서술된 설명과 함께, 본 발명의 원리를 설명하는 것을 돕는다. 가능한 어느 곳에서든, 같은 참조 숫자는 실시예의 같거나 또는 같은 요소들을 언급하기 위해서 도면 전체에 걸쳐서 사용되었다.
도 1은 벡터 지도(vector map)를 나타낸 도이다.
도 2는 융합 단백질의 SDS-PAGE 분석의 결과를 나타낸 도이다:
1 레인: RAP1-PE268 -313-E7-K3(환원된),
2 레인: 분자 마커,
3 레인: RAP1-PE268 -313-E7-K3(비-환원된).
도 3은 벡터 지도를 나타낸 도이다.
도 4는 본 발명의 한 실시예를 설명하는 개략도이다.
도 5a는 백신화(immunization) 스케쥴을 나타낸 도이다.
도 5b는 융합 단백질의 SDS-PAGE 분석의 결과를 나타낸 도이다:
1: RAP1-PE268 -313-E7-K3,
2: PE407-E7-K3,
M: 분자 질량 마커.
도 5c 및 d는 각각 다양한 융합 단백질 또는 플라시보(placebo)로 백신화된 동물 그룹에서 종양 크기 곡선 및 종양-없는 마우스의 백분율을 나타내는 도이다.
도 6는 융합 단백질의 준비를 위한 흐름 차트(왼쪽 패널), 및 융합 단백질의 SDS-PAGE 분석의 결과(오른쪽 패널)를 나타내는 도이다:
1, 4 레인: 분자 질량 마커,
2 레인: RAP1-CD28convPEt-E7-K3(마우스 CD28conv, 환원된),
3 레인: RAP1-CD28convPEt-E7-K3(마우스 CD28conv, 비-환원된),
5 레인: RAP1-CD28convPEt-E7-K3(인간 CD28conv, 환원된),
6 레인: RAP1-CD28convPEt-E7-K3(인간 CD28conv, 비-환원된).
도 7은 T 세포-민감성의 융합 단백질의 작용의 기전을 설명하는 개략도이다.
도 8은 다양한 종으로부터 CD28의 서열 정렬을 나타낸 도이다.
도 9a는 백신화 스케쥴을 나타낸 도이다.
도 9b 및 c는 각각 다양한 융합 단백질 또는 플라시보로 백신화된 동물 그룹에서 종양 크기 곡선 및 생존율을 나타낸 도이다.
도 10은 병원균으로부터의 DNA 삽입체(insert)를 포함하는 플라스미드(plasmid)를 생성하기 위한 RAP1-포함 벡터를 나타내는 개략도이다.
도 11은 다양한 병원균의 항원을 포함하는 융합 단백질의 제조를 설명하는 개략도이다.
도 12a 내지 f는 다양한 융합 단백질의 SDS-PAGE 분석의 결과를 나타내는 도이다.
도 13a 동물 그룹, 동물의 면역화에 사용되는 백신 및 투여용량(dosage)을 나타낸 도이다:
a: 각 도스는 PE407-DGD-K3(12 ㎍), PE407-M12-K3(6 ㎍), PE407-PQAB-K3(6 ㎍), 및 PE407-RSAB-K3(6 ㎍)의 혼합을 함유한다,
b: 각 도스는 RAP1-PE268 -313-DGD-K3(12 ㎍), RAP1-PE268 -313-M12-K3(6 ㎍), RAP1-PE268-313-PQAB-K3(6 ㎍), 및 RAP1-PE268 -313-RSAB-K3(6 ㎍)의 혼합을 함유한다,
c: 각 도스는 RAP1-CD28convPEt-DGD-K3(12 ㎍), RAP1-CD28convPEt-M12-K3(6 ㎍), RAP1-CD28convPEt-PQAB-K3(6 ㎍), 및 RAP1-CD28convPEt-RSAB-K3(6 ㎍)의 혼합을 함유한다.
도 13b는 면역화 스케쥴을 나타낸 도이다.
도 14 a 내지 d는 플라시보 또는 E716, E718, HCV중심 또는 HBx 항원을 포함하는 융합 단백질로 백신화된 도 13a의 동물 그룹으로부터 CD3+/CD4+ 비장세포(splenocyte) 및 CD3+/CD8+비장세포의 체외(ex vivo) 항원-특이적인 면역 반응 분석의 결과를 나타내는 표이다.
도 15a 내지 j는 플라시보 또는 PCV2(15a 및 b) 또는 PRRSV 항원(15c 내지 j)를 포함하는융합 단백질로 백신화된 13a의 동물 그룹으로부터 CD3+/CD8+ 비장세포(splenocyte) 및 CD3+/CD4+비장세포의 체외(ex vivo) 항원-특이적인 면역 반응 분석에서 IFNγ+세포 개수의 결과를 나타내는 도이다.
도 2는 융합 단백질의 SDS-PAGE 분석의 결과를 나타낸 도이다:
1 레인: RAP1-PE268 -313-E7-K3(환원된),
2 레인: 분자 마커,
3 레인: RAP1-PE268 -313-E7-K3(비-환원된).
도 3은 벡터 지도를 나타낸 도이다.
도 4는 본 발명의 한 실시예를 설명하는 개략도이다.
도 5a는 백신화(immunization) 스케쥴을 나타낸 도이다.
도 5b는 융합 단백질의 SDS-PAGE 분석의 결과를 나타낸 도이다:
1: RAP1-PE268 -313-E7-K3,
2: PE407-E7-K3,
M: 분자 질량 마커.
도 5c 및 d는 각각 다양한 융합 단백질 또는 플라시보(placebo)로 백신화된 동물 그룹에서 종양 크기 곡선 및 종양-없는 마우스의 백분율을 나타내는 도이다.
도 6는 융합 단백질의 준비를 위한 흐름 차트(왼쪽 패널), 및 융합 단백질의 SDS-PAGE 분석의 결과(오른쪽 패널)를 나타내는 도이다:
1, 4 레인: 분자 질량 마커,
2 레인: RAP1-CD28convPEt-E7-K3(마우스 CD28conv, 환원된),
3 레인: RAP1-CD28convPEt-E7-K3(마우스 CD28conv, 비-환원된),
5 레인: RAP1-CD28convPEt-E7-K3(인간 CD28conv, 환원된),
6 레인: RAP1-CD28convPEt-E7-K3(인간 CD28conv, 비-환원된).
도 7은 T 세포-민감성의 융합 단백질의 작용의 기전을 설명하는 개략도이다.
도 8은 다양한 종으로부터 CD28의 서열 정렬을 나타낸 도이다.
도 9a는 백신화 스케쥴을 나타낸 도이다.
도 9b 및 c는 각각 다양한 융합 단백질 또는 플라시보로 백신화된 동물 그룹에서 종양 크기 곡선 및 생존율을 나타낸 도이다.
도 10은 병원균으로부터의 DNA 삽입체(insert)를 포함하는 플라스미드(plasmid)를 생성하기 위한 RAP1-포함 벡터를 나타내는 개략도이다.
도 11은 다양한 병원균의 항원을 포함하는 융합 단백질의 제조를 설명하는 개략도이다.
도 12a 내지 f는 다양한 융합 단백질의 SDS-PAGE 분석의 결과를 나타내는 도이다.
도 13a 동물 그룹, 동물의 면역화에 사용되는 백신 및 투여용량(dosage)을 나타낸 도이다:
a: 각 도스는 PE407-DGD-K3(12 ㎍), PE407-M12-K3(6 ㎍), PE407-PQAB-K3(6 ㎍), 및 PE407-RSAB-K3(6 ㎍)의 혼합을 함유한다,
b: 각 도스는 RAP1-PE268 -313-DGD-K3(12 ㎍), RAP1-PE268 -313-M12-K3(6 ㎍), RAP1-PE268-313-PQAB-K3(6 ㎍), 및 RAP1-PE268 -313-RSAB-K3(6 ㎍)의 혼합을 함유한다,
c: 각 도스는 RAP1-CD28convPEt-DGD-K3(12 ㎍), RAP1-CD28convPEt-M12-K3(6 ㎍), RAP1-CD28convPEt-PQAB-K3(6 ㎍), 및 RAP1-CD28convPEt-RSAB-K3(6 ㎍)의 혼합을 함유한다.
도 13b는 면역화 스케쥴을 나타낸 도이다.
도 14 a 내지 d는 플라시보 또는 E716, E718, HCV중심 또는 HBx 항원을 포함하는 융합 단백질로 백신화된 도 13a의 동물 그룹으로부터 CD3+/CD4+ 비장세포(splenocyte) 및 CD3+/CD8+비장세포의 체외(ex vivo) 항원-특이적인 면역 반응 분석의 결과를 나타내는 표이다.
도 15a 내지 j는 플라시보 또는 PCV2(15a 및 b) 또는 PRRSV 항원(15c 내지 j)를 포함하는융합 단백질로 백신화된 13a의 동물 그룹으로부터 CD3+/CD8+ 비장세포(splenocyte) 및 CD3+/CD4+비장세포의 체외(ex vivo) 항원-특이적인 면역 반응 분석에서 IFNγ+세포 개수의 결과를 나타내는 도이다.
본 발명은 오직 당업계에 있는 기술자에게는 자명한 수많은 변형 및 변이 기술로서 설명되는 것으로 의도되는 하기의 실험예에서 더 명확하게 기술된다. 본 발명의 다양한 실시예를 이제 자세하게 기술한다. 도면을 참조하는 것은, 전체에 걸쳐서 숫자가 같은 것이 구성요소 같은 것을 가르키는 것이다. 본 명세서의 기술 및 청구항 전체에 사용된 하기의, "단수어(a)", "단수어(an)", 및 "한정어(the)"는 문맥상 명백하게 다른 것을 나타내지 않는다면 복수 참조를 포함하는 의미이다. 또한, 본 명세서의 기술 및 청구항 전체에서 사용된 하기의, "안에(in)"는 문맥상 명백하게 다른 것을 나타내지 않는다면 "안(in)" 및 "위(on)"을 포함하는 의미이다. 더 나아가, 본 명세서의 제목 및 부제목은 본 발명의 범위에 영향을 주지 않고, 독자의 편의를 위해서 사용하였다. 추가적으로, 본 명세서에 사용된 몇몇의 용어는 하기에 더 자세하게 정의하였다.
정의
본 명세서에서 사용된 용어는 일반적으로 본 발명의 문맥 안에서, 당업계에서 및 각 용어가 사용된 특정 문맥에서의 그들의 보통의 의미를 갖는다. 본 발명을 설명하는데 사용된 특정 용어는 본 발명의 기술을 고려하는 전문가에게 추가적인 지도를 제공하기 위해 하기 또는 본 명세서의 어딘가에서 논의하였다. 편의를 위해서, 특정 용어는, 예를 들면 이탤릭체(italics) 및/또는 따옴표(quotation marks)의 형태로 강조하였다. 같은 것이 하나 이상의 방법으로 얘기할 수 있다는 것이 이해될 것이다. 결론적으로, 대체적인 언어 및 동의어는 본 명세서에서 논의되는 한 또는 그 이상의 용어로 사용되며, 용어가 본 명세서에서 확대되거나 논의되던지 간에 어떠한 특별한 중요성을 갖지 않는다. 특정 용어에 대한 동의어는 제공하였다. 하나 또는 이상의 동의어의 설명이 다른 동의어의 사용을 배재 하는 것은 아니다. 본 명세서에서 논의된 임의의 용어의 예시를 포함하는 본 명세서의 어딘가에 있는 실험예의 사용은 오직 설명적이며, 어떠한 방식으로든 본 발명 및 임의의 예시된 용어의 범위 및 의미를 제한한다. 이처럼, 본 발명은 본 명세서에 제시된 다양한 실시예에 한정되지 않는다.
정의되지 않는 한, 본 명세서에 사용된 모든 기술적 및 과학적인 용어들은 본 발명이 적용되는 당업계의 평범한 기술을 가진 자에 의해 일반적으로 이해되는 것과 같은 의미를 가진다. 논란의 경우에, 용어를 포함하는 본 문서는 조절될 것이다.
용어 "항원-제시 세포(APC) 또는 부수적인 세포(accessory cell)"은 세포 표면에 주요 조직적합성 복합체(major histocompatibility complexes; MHCs)와 복합체를 이루는 외부 항원을 나타내는 세포를 말한다. T-세포는 T-세포 수용체(T-cell receptors; TCRs)를 이용하여 이러한 복합체들을 인식할 것이다. 이러한 세포는 항원을 처리하며 이들을 T-세포로 제시한다. 다양한 종류의 전문적인 항원-제시 세포는 수지상 세포(dendritic cells; DCs), 대식세포(macrophages), 단핵 백혈구(monocytes), 및 특정 B-세포이다.
용어 "항원-제시 세포(APC)-결합 도메인"은 항원-제시 세포(APC)에 결합할 수 있는 도메인(폴리펩타이드 인 것)을 말한다. 상기 APC-결합 도메인은 서열번호 5, 6, 7, 8, 및 9로 구성된 군으로부터 선별된 서열과 최소 90% 동일한 아미노산 서열을 포함하는 폴리펩타이드 일 수 있다. APC-결합 도메인은 APC에 있는 수용체를 인식하고 결합하는 리간드(ligand)이다.
분화 91의 집단(cluster of differentiation 91; CD91)은 세포의 막에서 수용체를 구성하고 수용체-매개의 세포내섭취에 연관된 단백질이다.
용어 "단백질 형질도입(transduction) 도메인"은 T-세포에 민감하여 항원-특이적인 T 세포 반응을 강화할 수 있고, 및/ 또한 항원 제시의 클래스 I 주요 조직적합성 복합체(class I major histocompatibility complex; MHC-I) 경로(즉, 세포독성의 T 세포 경로)로 항원을 가이드 또는 직접 향하도록 하는 기능을 갖고 있는 폴리펩타이드 또는 융합 폴리펩타이드를 말한다.
용어 "T 세포에 민감하게"는 일반적으로 CD8+ 및 CD4+ T 세포가 민감하고 결로적으로, 항원 공격(challenge)에 대한 CD8+(CTL) 및 CD4+ T 세포 반응이 강화된 것을 말한다. 항원-특이적인 세포 매개의 면역 반응은 항원에 대한 반응에서 항원-특이적으로 유도된 γ-인터페론(interferon)의 생산을 정량화하여 측정하였다. 예를 들면, 민감성 신호가 없을 때(즉, 단백질 형질도입 도메인 없이), 항원 단독은 약하거나 세포 매개의 면역 반응을 전혀 유도하지 못하는데, 즉 CD8+ 및 CD4+ T 세포로부터 항원-특이적인 γ-인터페론의 생산이 약하거나 없는데, 반면 민감성 신호(단백질 형질도입 도메인)의 존재에서는, 항원이 강화된 세포 매개의 면역 반응을 유도할 수 있다. 그러므로, 민감성 신호(단백질 형질도입 도메인)의 기능은 숙주 안에서 CD8+ 및 CD4+ T 세포에 민감한 것이고 이를 통해 이후에 숙주가 항원에 의해 공격되었을 때, 항원이 이전의 CD8+ 및 CD4+ T 세포 민감화로 인해 강화된 항원-특이적인 세포 매개의 면역 반응을 유도할 수 있을 것이다.
단백질 형질도입 도메인은 하기로 구성된 그룹으로부터 선택된 펩타이드 및/또는 폴리펩타이드이다:
(i) T 세포 민감성 신호-형질도입 펩타이드, 링커(linker) 및 전좌 펩타이드로 구성된 융합 폴리펩타이드, 여기서:
(1) 상기 T 세포 민감성 신호-형질도입 펩타이드는 융합 폴리펩타이드의 N-말단에 위치한 것이다;
(2) 상기 링커는 T 세포 민감성 신호-형질도입 펩타이드 및 전좌 펩타이드를 연결하는, 서열번호 15를 포함한다; 및
(3) 상기 전좌 펩타이드는 34 내지 112 아미노산 잔기의 길이이며 서열번호 3, 20 및 4와 최소 90% 동일한 아미노산 서열을 포함한다;
(ii) T 세포-민감성 신호-형질도입 펩타이드; 및
(iii) 서열번호 3, 20 및 4와 최소 90% 동일한 아미노산 서열을 포함하는 34 내지 112 아미노산 잔기의 길이의 전좌 펩타이드.
단백질 형질도입 도메인은 T 세포 민감성 신호-형질도입 펩타이드, 링커, 및 전좌 펩타이드를 포함하는 융합 폴리펩타이드인 "융합 폴리펩타이드"일 수 있다. 예를 들면, 상기 융합 폴리펩타이드는 "CD28convPEt"일 수 있다.
용어 "CD28conv"는 "T 세포 민감성 신호-형질도입 펩타이드"인 CD28 보존된 영역(conserved region)을 말한다. 이는 CD28에 대항하는 항체를 유도하기 위한 항원결정기(epitope)이다.
용어 "PEt"는 34 내지 112 아미노산 잔기 길이의 전좌를 말한다.
링커는 "CD28conv" 및 "PEt"의 사이에 존재한다. 상기 융합 폴리펩타이드 "CD28convPEt"의 방향 또는 정렬은 "CD28conv"(또는 T 세포 민감성 신호-형질도입 펩타이드)가 반드시 PEt의 상위(upstream)에 존재해야만 하며, 즉 PEt는 강화된 T-세포 반응을 얻기 위해 반드시 "CD28conv"의 C-말단에 위치해야한다는 점에서 중요하다. 상기 "CD28convPEt"는 역방향의 융합 펩타이드 PEtCD28conv보다 CD28conv에 대한 훨씬 높게 IgG 역가(CD28-특이적인 길항(agonist) 항체)를 올릴 수 있다. 상기 CD28-특이적인 길항 항체는 CD4+ 및 CD8+ T 세포 모두에 민감할 수 있다. 옳은 방향의 융합 폴리펩타이드 CD28convPEt는 CD28conv 및 PEt 도메인 사이의 링커(RXRXKR)을 포함한다. 상기 링커는 항원 제시 세포(APC) 특이적인 프로테아제(protease)(카뎁신 L(cathepsin L)) 절단 위치 Lys-Arg(KR)을 포함한다. 그러므로, 융합 단백질 RAP1-CD28convPEt-항원-K3는 하기의 두 개의 절편으로 잘릴 수 있다: RAP1-CD28conv 및 PEt-항원-K3. RAP1-CD28conv 절편은 리소좀에 의해 더 잘려질 수 있으며 CD28conv의 항원결정기는 CD28 길항 항체를 생산하는 체액성 면역 반응을 유도하는, MHC II 경로를 통해 APC 세포 표면으로 제시된다. 그러므로, CD28 길항 항체는 B 세포에 의해 생산된다. 상기 CD28 길항 항체는 T 세포 표면의 CD28에 결합 및 T 세포(CD4+ 및 CD8+ T 세포)를 전-활성화(pre-activate)할 수 있다.
"T 세포-민감성 신호-형질도입 펩타이드"는 28 내지 53 아미노산 잔기 길이이고 Xaa8은 I 또는 L; Xaa10은 V, F 또는 A; Xaa11은 M 또는 L; Mxx17은 L 또는 I인 서열번호 31과 최소 90% 동일한 아미노산 서열을 포함한다.
상기 T 세포-민감성 신호-형질도입 펩타이드는 (X)2는 I 또는 L; (X)4는 V, F 또는 A; (X)5는 M 또는 L인 K1(I/L)2E3(V/F/A)4(M/L)5Y6P7P8P9Y10(서열번호 32)의 중요 지역을 포함한다.
마우스에 특이적인 T 세포 민감성 신호-형질도입 펩타이드(TDIYFCKIEFMYPPPYLDNEKSNGTIIH; 서열번호 31, 여기서 X8은 I, X10은 F, X11은 M)는 하기의 실험예에 사용되었다.
도 7은 예시로서 융합 단백질 RAP1-CD28convPEt-E7-K3를 사용하는 T 세포-민감성 융합 단백질의 작용의 기전을 설명하는 개략도이다. RAP1-CD28convPEt-E7-K3는 N-말단에서 C-말단으로 다음을 포함한다: (1) N-말단에서 전장 RAP1의 도메인 III, (2) CD28conv, (3) 링커, (4) 수도모나스 엑소톡신 A(Pseudomonas Exotoxin A)로부터의 변형된 전좌 펩타이드, (5) 전장 HPV 타입 16 E7 단백질 및 (6) C-말단에서 ER 유지 신호로서 세 개의 KDEL. 상기 HPV16E7 단백질은 융합 단백질과 함께 면역화된 개체의 세포 안에 있는 항원결정기로 진행된다. 상기 RAP1-CD28convPEt-E7-K3 단백질은 APC(수지상 세포 같은) 흡수(uptake) 효율을 개선하고 프로테아좀(proteasome) 경로를 향한 후, MHC I 복합체를 통해 제시되도록 HPV16E7 항원 진행의 강화할 수 있도록 설계되었다. 백신 RAP1-CD28convPEt-E7-K3에 의해 유도되는 HPV16E7 단백질-특이적인 CTL 면역 반응의 작요의 기전은 도 7에 나타나있다: (a) 백신은 APC(수지상 세포와 같은) 표면 수용체(CD91)에 결합하고 세포내섭취에 의해 내재화된다(internalized); (b1) RAP1-CD28convPEt-E7-K3는 전좌 펩타이드 PEt의 전 위치에서 카뎁신 L 프로테아제 소화에 의해 단백질 가수분해(proteolytic hydrolysis)가 진행된다; (b2) 또는 E.R.로 재활용되거나 전좌 펩타이드 PEt의 전 위치에서 퓨린(furin) 프로테아제에 의해 단백질 가수분해가 진행된다; (b3) 동시에, RAP1-CD28conv는 리소조말(lysosomal) 프로테아제에 의해 조화되고 CD28conv의 항원결정기는 MHC II를 통해 세포 표면에 제시되고 T 세포를 전-활성화 할 수 있는 CD28 길항 항체 생산을 유도한다; (c) 가장 중요한 단계는 전좌 펩타이드(PEt)에 의해 리소좀으로부터 세포질성 부위(cytoplasmic compartment)로 PEt-E7-K3의 막관통성(transmembrane) 전좌이다; (d) PEt-E7-K3는 프로테아좀 경로를 통해 소화가 지행되고, E7의 항원결정기는 MHC I 복합체에 의해 제시되며 E7-특이적인 세포 매개의 면역 반응을 유도한다.
도 8은 다양한 종으로부터 CD28 보존 영역의 서열 정렬 및 일치 서열을 나타낸다. 일치 서열 내의 밑줄 친 서열(KIEVMYPPPY; 서열번호 32, 여기서 X2는 I, X4는V, X5는 M)은 CD28 길항 항체 인식 및 결합을 위한 주요 영역이다. 상기 주요 영역 서열은 K1(I/L)2E3(V/F/A)4(M/L)5Y6P7P8P9Y10로 나타낼 수 있고, 여기서 오직 네번째 아미노산 잔기가 종-특이적이고 V는 인간, 생쥐, 돼지, 소, 양, 개 및 말; F는 마우스, 및 V는 칠면조이다. 상기 주요 영역 서열은 K1(X)2E3(X)4(X)5Y6P7P8P9Y10(서열번호 32)로서 나타낼 수 있고, 여기서 (X)2는 I 또는 L; (X)4는 V, F 또는 A, (X)5는 M 또는 L이다.
PEt는 서열번호 3 또는 20과 최소 90% 동일한 아미노산 서열을 포함할 수 있다. 예를 들면, PEt의 아미노산 서열은 PE의 a.a.280 내지 a.a.313(서열번호 3), a.a.268 내지 a.a.313(서열번호 20), a.a.253 내지 a.a.313, 또는 a.a.253 내지 a.a.364(서열번호 4)일 수 있다. 즉, PEt의 아미노산 서열은 a.a.280 내지 a.a.313(서열번호 3) 필수적인 절편을 포함하는 것만큼 긴 PE 도메인 II(a.a.253 내지 a.a.364; 서열번호 4)의 임의의 영역을 포함할 수 있다.
항원은 형원균에 의해 야기되는 질명에 책임이 있거나, 또는 병원균에 의해 감염된 숙주 내에서 면역적인 반응을 유도할 수 있는 병원균 단백질, 폴리펩타이드 또는 펩타이드, 또는 종양 세포에서 특이적으로 발현되는 폴리펩타이드인 종양-연관된 항원(TAA)일 수 있다. 상기 항원은 인간 파필로마바이러스(Human Papillomavirus; HPV), PRRSV, HIV-1, 댕기 바이러스(Dangue virus), C형 간염 바이러스(Hepatitis C virus; HCV), B형 간염 바이러스(Hepatitis B virus; HBV), 돼지 서코바이러스(Porcine Circovirus 2; PCV2), 비소세포성 폐암(non-small cell lung cancer), 유방암(breast carcinoma), 흑색종(melanoma), 림프종(lymphomas), 대장암(colon carcinoma), 간세포 암종(hepatocellular carcinoma) 및 이러한 것들의 임의의 조합을 포함하는 병원균 또는 암 세포로부터 선별될 수 있으나 이에 한정되지 않는다. 상기 항원은 하나 또는 그 이상의 병원성 단백질로부터 선택된 둘 또는 그 이상의 항원의 융합으로부터의 융합 항원일 수 있다. 예를 들면, PRRSV ORF6 및 ORF5 절편의 융합 항원, 또는 PRRSV 및 PCV2 병원균으로부터 항원성 단백질의 융합이다.
골지체 유지 서열의 기능은 세포유입성 부분으로부터 ER로의 항원의 전좌를 돕고 루멘(lumen)에서 이를 유지하도록 한다. 이는 Lys Asp Glu Leu(KDEL) 또는 RDEL의 서열을 포함한다. ER 서열은 KKDLRDELKDEL (서열번호 16), KKDELRDELKDEL (서열번호 17), KKDELRVELKDEL (서열번호 18)의 서열을 포함하거나, 또는 필수적으로 구성되거나, 또는 구성된다.
39 kDa의 분자량을 갖는 수용체-연관 단백질(RAP1)은 ER에 존재하는 단백질 및 LDL 수용체-연관 단백질을 위한 분자 샤페론(chaperone)이다. 이는 CD91(Kd ~ 3 nM)에 대해 높은 결합 친화력(affinity)를 갖고 있으며 세 개의 기능적-유사성 도메인으로 구성되었다.
본 발명은 본 발명에 따르는 융합 다백질에 의한 T 세포 매개의 면역 반응의 유도 및 강화의 발견과 연관되어있다. RAP1-CD28convPEt-E7-K3를 사용하는 예시에서, RAP1-CD28convPEt-E7-K3 백신의 전략은 표적 항체 E7을 나타내는 HPV16 감염된 세포를 인식할 수 있는 T 세포의 생산 및 활성을 자극하는 것에 중점적으로 집중되어있다. 항체를 수지상 세포로 전달에 의해, 이는 항원-특이적인 CD8+ T 세포 및 CD4+ T 세포를 생성할 수 있다. 타입 1-조력자(helper) CD4+ T 세포는 특히 세포독성 CD8+ T 세포의 면역 반응을 효과적으로 자극하고 증가시킬 수 있다. 종합하여, 이러한 적응 면역 시스템의 두 개의 팔은 정상 조직에 현저한 피해를 주지 않으며 신체의 다양한 위치에서 HPV16-감염 세포 또는 HPV16-연관된 종양 세포를 사멸할 수 있는 특이성 및 효능을 갖고 있다.
용어 "개체"는 인간 또는 비-인간 동물을 말한다.
용어 "치료(treating)" 또는 "치료(treatment)"는 암 또는 감염을 갖거나, 이러한 질병에 대해 증상 또는 소질(predisposition)을 갖고 있는 개체에 질병의 예방, 증상 또는 그것에 대한 소질을 예방하거나 치료, 완화(alleviate), 안정, 요법(remedy), 개선의 목적을 갖고, 효과적인 양의 융합 단백질을 투여하는 것을 말한다. 이러한 개체는 어떠한 적합한 진단 방법에 의한 결론에 기초하는 건강 관리 전문가에 의해 확인될 수 있다.
용어 "효과적인 양"은 치료된 개체에서 치료적인 효과를 부여하는데에 필요한 활성화된 화합물(active compound)의 양을 말한다. 효과적인 도스(dose)는 다향할 것이고, 투여의 경로, 첨가제(excipient) 사용, 및 다른 치료적 치료와의 공동-사용의 가능성에 의존하여 당업계의 기술자에 의해 인식될 수 있다.
약어: CD28, 분화의 집단(Cluster of Differentiation) 28.
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본 발명, 예시적 기기, 기구, 방법 및 본 발명의 실시예를 따르는 상기와 연관된 결과들의 범위를 한정하는 의도가 없이 하기에 제공하였다. 제목 또는 부체목은 독자의 편의를 위해 실험예에서 사용되었는데, 본 발명의 범위를 한정하는 방법이 아닌 것을 알린다. 게다가, 특정 이론은 본 명세서에서 제안되고 및 기술되었다; 그러나, 상기가 옳고 그름에 관계없이, 본 발명은 임의의 특정 이론이나 작용 방식에 관계없이 본 발명에 따라 실시되는 대로 길게 발명의 범위를 제한한다.
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1> 발현 벡터의 제조
도 1은 RAP1 도메인 3(서열번호 5), 전좌 최소 필수적 펩타이드(PE268 -313;서열번호 20), 항원 E7, 및 골지체 유지 서열(K3, 또는 KDEL 신호; 서열번호 16, 17, 18, 또는 19)를 포함하는 융합 단백질 PERAP1-PE268 -313-E7-K3의 발현 벡터를 나타낸다. 전좌 최소 필수적 펩타이드(PE268-313;서열번호 20)은 a.a.268 내지 a.a.313의 PE(서열번호 10) 폴리펩타이드 서열 영역으로부터 얻었다.
상기 발현 벡터는 도 10에 나타난 플라스미드 RAP1-K3를 사용하여 제조하였다. RAP-1 도메인 3 및 K3의 융합 유전자를 포함하는 상기 플라스미드 RAP1-K3(도 10)는 하기와 같이 생성하였다: NdeIRAP1-(EcoRI, XhoI )-K3XhoI 을 암호화하는 DNA 절편을 PCR 방법으로 합성하고 플라스미드 RAP1-K3(도 10)을 얻기 위한 가나마이신 저항 유전자를 갖고 있는 플라스미드 pUC18 백본에 접합하였다. 발현 벡터 RAP1-PE268-313-E7-K3(도 1)을 생성하기 위해, PE268 -313-E7을 암호화하는 DNA 절편을 플라스미드 RAP1-K3(도 10)에 삽입하였다. 비슷한 방법을 사용하여, 다양한 융합 펩타이드를 암호화하는 DNA 절편을 PCR로 생성하고(도 11) 다양한 융합 단백질(도 12a 내지 f)에 대한 발현 벡터를 생성하기 위해 플라스미드 RAP1-K3(도 10)로 각각 삽입하였다. 도 3은 융합 단백질 RAP1-CD28convPEt-E7-K3에 대한 발현 벡터를 나타낸다(도 3).
절편 RAP1-CD28convPEt는 카뎁신 L 및 퓨린 프로테아제 절단 자리를 갖고 있다. 도 4는 절편 RAP1-CD28convPEt의 중요성을 설명하기 위한 융합 단백질 RAP1-CD28convPEt-E7-K3의 아미노산 서열 지도를 나타낸다. 두 개의 화살표는 각각 카뎁신 L 및 퓨린 프로테아제 절단 위치를 표시한다. 상기 두 개의 절단 위치는 CD28conv 및 PEt-E7의 접합을 위한 링커로서 작용할 뿐만 아니라 절편 PEt-E7-K3를 리소좀으로부터 세포질로 용리되도록 융합 단백질의 절단을 허용한다. 다양한 병원균으로부터 어떠한 다른 흥미로운 항원들은 CD28convPEt와 융합하기 위해 E7과 대체할 수 있으며 상기 융합 생성물은 도 11에 기술된 융합 단백질을 위한 다양한 종류의 발현 벡터를 만들기 위해 도 10의 플라스미드로 삽입될 수 있다.
PE268 -313의 서열은: pletftrhrqprgweqleqcgypvqrlvalylaarlswnqvdqvir(서열번호 20)이다. CD28convPEt의 전체 서열은 하기와 같다: tdiyfckiefmypppyldneksngtiihrarykrgweqleqcgypvqrlvalylaarlswnqvdqvirgs(서열번호 30), 밑줄 친 서열은 카뎁신 L 및 퓨린 프로테아제 절단 위치를 포함하는 링커 서열을 나타낸다.
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2> 단백질 발현
단백질 발현 벡터를 갖고 있는 E. coli BL 21 세포는 37℃의 25 ㎍/ml 가나마이신을 함유하는 루리아 버타니 브로스(Luria Bertani broth)에서 배양하였다. 배양액이 초기 로그 단계에 도달하였을 때, (A600=0.1 내지 0.4), 유도(induction)를 위해 최종 농도 0.5 내지 2 mM의 이소프로필-1-싸이오-β-D-갈락토피라노시드(isopropyl-1-thio-β-D-galactopyranoside; IPTG)를 첨가하였다. IPTG 유도 4시간 후에 세포를 수확하였고 초음파분쇄(sonication)를 이용하여 세포를 분쇄하였다. 상기 과발현 단백질-함유 내포체(inclusion body)를 분리하여 8 M 우레아(urea)/TN 완충용액(8 M 우레아, 50 mM 트리스, 50 mM NaCl, pH 8.0)에 용해하였다.
융합 단백질 RAP1-CD28convPEt-E7-K3의 재접힘은 4℃에서 하루 동안 TNZ 완충용액(50 mM 트리스, 50 mM NaCl 및 0.01 mM ZnCl2, pH 8.0)의 50X 부피에 대항하는 투석(dialysis)으로 수행하였다. 상기 재접힘된 단백질은 환원(디싸이오쓰게이톨(dithiothreitol와 함께; +DTT) 및 비-환원(-DTT) 조건에서 SDS-PAGE로 수행하였다(도 2). 상기 결과는 비-환원 조건에서 대부분의 재접힘된 단백질이 모노머(monomers)였는데, 이는 RAP1 융합 단백질이 재접힘이 쉽고 응집되지 않는다는 것을 나타낸다(도 2).
도 6a는 E. coli 세포의 내포체로부터 발현되고 추출된 융합 단백질 RAP1-CD28convPEt-E7-K3(마우스 CD28conv 또는 인간 CD28conv)를 설명하는 흐름 도이다. SDS-PAGE 분석은 융합 단백질이 재접힘이 잘되었음 나타낸다(도 6b).
도 11은 상기에 기술된 비슷한 방법을 사용하여 발현된 융합 단백질의 목록을 나타낸다: (1) RAP1-PE268 -313-E7-K3; (2) RAP1-CD28convPEt-E7-K3; (3) RAP1-CD28PEt-E718-K3; (4) RAP1-HCVcore-K3; (5) RAP1-CD28conv-HCVcore-K3; (6) RAP1-CD28convPEt-HCVcore-K3; (7) RAP1-HBx; (8) RAP1-HBx-K3; (9) RAP1-CD28conv-HBx; (10) RAP1-CD28conv-HBx-K3; (11) RAP1-CD28convPEt-HBx; (12) RAP1-CD28convPEt-HBx-K3; (13) RAP1-PCV2ORF2-K3; (14) RAP1- PE268 -313-PCV2ORF2-K3; (15) RAP1-CD28convPEt-PCV2ORF2-K3; (16) RAP1-PE268 -313-DGD-K3; (17) RAP1-PE268 -313-M12-K3; (18) RAP1-PE268 -313-PQAB-K3; (19) RAP1-PE268 -313-RSAB-K3; (20) RAP1-CD28convPEt-DGD-K3; (21) RAP1-CD28convPEt-M12-K3; (22) RAP1-CD28convPEt-PQAB-K3; (23) RAP1-CD28convPEt-RSAB-K3. 이러한 융합 단백질들은 상기에 기술 것과 같은 방법을 사용하여 재접힘되었다. SDS-PAGE의 결과는 이러한 융합 단백질들이 모두 잘 재접힘되어 백신의 준비를 위해 사용됨을 나타낸다(도 12a 내지 f).
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3>
RAP1
-
PE
268
-313
-
E7
-
K3
는 인간 파필로마 바이러스(
HPV
) 타입 16
E7
단백질에 의해 유도된 종양의 성장을 억제함
PE407-E7-K3 및 RAP1-PE268 -313-E7-K3의 융합 단백질은 상기와 같이 발현되었고 SDS-PAGE에 의해 단백질 재접힘을 검사하였다(도 5b). 마우스는 HPV-16 타입 암종(carcinoma)을 유도하기 위해 피하 주사(s.c. injection)를 통해 2×103 TC-01 세포(HPV 타입 16 E7 유전자를 갖는 마우스 폐 표피 세포 주)로 시도하였다. TC-01 세포 시도 12일 후에, 마우스는 3주 동안 한 주에 한번 씩, 아주번트로서 AS04C(GlaxoSmithKline)과 함께 플라시보(PBS+알루미늄 포스페이트), PE407-E7-K3(200 ㎍/도스) 또는 RAP1-PE268 -313-E7-K3(200 ㎍/도스)를 피하주사로 백신화하였다(도 5a). 세포독성 T 림프구(lymphocyte)-강화 아주번트인 AS04C는 MPL(모노포스포릴 지방 A(monophosphoryl lipid A), 면역 강화제(potentiator)) 및 알루미늄 포스페이트(항원 전달을 위한 단백질 흡습제)를 포함한다. 용어 "K3"는 아미노산 서열 KDELKDELKDEL(서열번호 19)를 의미한다. 각 그룹에서 종양의 크기 및 종양-없는 동물의 숫자는 기록되었다(도 5c 및 d). 종양의 성장은 아주번트로서 AS04C와 함께 두 개의 백신 PE407-E7-K3 및 RAP1-PE268 -313-E7-K3 모두에 의해서 현저하게 억제되었다. 반면, RAP1-PE268 -313-E7-K3로 백신화된 마우스 그룹이 더 높은 종양-없는 마우스의 비율을 나타냈다. 상기는 백신 RAP1-PE268 -313-E7-K3가 종양의 성장을 억제하는 면에서 PE407-E7-K3만큼 효과적이거나 또는 더 낫고 종양-없는 동물의 백분율의 증가면에서 더 나은 것을 나타낸다.
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4>
RAP1
-
CD28convPE
t
-
E7
-
K3
가 인간 파필로마 바이러스(
HPV
) 타입 16 E7 단백질에 의해 유도되는 종양의 성장을 억제하고 생존율을 증가시킴
융합 단백질 PE407-E7-K3 및 RAP1-CD28convPEt-E7-K3와 면역 강화제의 첨가 또는 무첨가의 종양 크기 및 생존율에 대한 효과를 관찰하였다. 마우스는 높은 도스의 TC-01 세포(3×104)를 피하 주사로 시도되었다. 시도 7일 후, 마우스는 3주 동안 한 주에 한번 씩 면역 강화제 GPI-0100 또는 단백질 흡습제 알루미늄 포스페이트와 함께 플라시보, PE407-E7-K3(100 ㎍/도스) 또는 RAP1-CD28convPEt-E7-K3(100 ㎍/도스)를 피하주사로 백신화하였다(도 9). GPI-0100은 Th1/CTL 자극 아주번트(면역 강화제)이다. 각 그룹에서 종양의 크기 및 생존율을 기록하였다(도 9b 및 c). 아주번트 GPI-0100과 결합하였을 때, PE407-E7-K3 및 RAP1-CD28convPEt-E7-K3 모두 종양 성장을 억제하였다. 예상외로, RAP1-CD28convPEt-E7-K3의 종양 성장 억제에 대한 효과가 아주번트에 의존하지 않는다는 것을 발견하였다. 아주번트 GPI-0100 보다 흡습제 알루미늄 포스페이트에 결합하였을 때, RAP1-CD28convPEt-E7-K3는 면역 강화제 GPI-0100과 결합되었을 때와 같은 효능으로 종양의 성장을 현저하게 억제하였다(도 9b, 색칠 되지 않은 삼각형 대 색칠 된 역 삼각형).
반면, PE407-E7-K3의 종양 성장 억제에 대한 효능은 아주번트에 의존하였다. 흡습제 알루미늄 포스페이트와 결합하였을 때, PE407-E7-K3는 면역 강화제 GPI-0100과 결합하였을 때보다 효능이 덜 하였다(도 9b, 색칠 된 사각형 대 색칠 되지 않은 원형).
한편, 면역 강화제 GPI-0100 또는 흡습체 알루미늄 포스페이트와 결합된 RAP1-CD28convPEt-E7-K3를 투여한 마우스는 GPI-0100 또는 알루미늄 포스페이트와 결합된 PE407-E7-K3로 백신화한 그룹보다 더 좋은 생존율을 나타냈다(도 9c). 이는 RAP1-CD28convPEt-E7-K3가 면역 강화제 GPI-0100 없이도 Th1/CTL 면역 반응을 유도할 수 있음을 나타낸다. 상기 결과는 또한 융합 단백질 RAP1-CD28convPEt-E7-K3가 동물의 생존율을 높이기 위한 백신으로서 PE407-E7-K3보다 더 뛰어남을 나타낸다.
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5> 면역원성(
immunogenicity
) 분석
다양한 백신의 면역원성을 시험하였다. 간단하게, 마우스는 하기의 그룹으로 나누었다: HPV16 E7, HPV 18 E7, HCV 중심, HBV HBx, PCV2 ORF2 및 PRRSV(도 13a). 각 그룹은 부그룹(subgroups)으로 더 나누었고, 각 부그룹은 3주 동안 한 주에 한번 씩 피하 주사를 통해 플라시보 또는 병원균의 특정 항원, 또는 특정 항원들을 표적하도록 설계된 백신으로 주사하였다(도 13b). PRRSV를 표적하는 백신을 제외하고, 각 백신은 병원균의 최소 하나의 항원을 포함하는 단일 융합 단백질인, 당일 융합 단백질 및 흡습제 알루미늄 포스페이트로 구성되었다. 상기 항원은 병원균으로부터 전장 단백질이거나 또는 병원균의 항원의 최소 하나의 항원결정기를 포함하는 비-전장 단백질, 또는 병원균의 서로 다른 단백질로부터 선별된, 두 개 또는 그 이상의 항원의 융합 펩타이드이다.
백신화 스케쥴은, 도 13a 및 b에 백신 및 도스(dose)를 설명하였다. 간단하게, 마우스는 도 13a에 나열된 백신으로 3주 동안 한 주에 한번 씩 백신화하였다. 모든 마우스는 마지막 면역화 7일 후에 희생되었으며, 비장(spleen)을 수확하였다. 비장세포(splenocytes)를 분리한 후, 1 ㎍/ml의 GolgiPlug(BD Pharmingen, San Diego, CA)의 존재하에서 비장세포를 자극할 수 있는 병원균의 개별적인 재조합 항원 10 ㎍/ml과 함께 6 웰 플레이트(2×107 세포/2 ml/웰)에서 37℃, 16시간 동안 배양하였다.
자극된 비장세포는 FACScan 완충용액으로 세척하고 세포 표면 마커 CD8a, CD4, 및 CD3는 피코에리쓰린(phycoerythrin)-결합된 단일클론 생쥐 항-마우스 CD8a, AF700-결합된 단일항체 생쥐 항-마우스 CD4 및 AF647-결합된 단일클론 생쥐 항-마우스 CD3 항체로 염색되었다. 상기 세포는 투과성화(permeabilized)하였고 제조자의 지시(BD Pharmingen)에 따라 Cytofix/Cytoperm 키트에 의해 고정화하였다. 세포내(intracellular) IFN-γ는 면역 반응 및 사이토카인(cytokine) 수준을 측정하기 위하여 AF488-결합 생쥐 항-마우스 IFN-γ로 염색하였다. 유동 세포 분석법(flow cytometry)은 Kaluza 분석 소프트웨어(Beckman Coulter)와 함께 Gallios 유동 세포 분석법을 사용하여 수행하였다.
하기 PRRSV 백신을 면역원성에 대해 시험하였다: PE407-PRRSV-K3, RAP1-PE268 -313-PRRSV-K3 또는 RAP1-CD28convPEt-PRRSV-K3 백신. 각 백신은 네 개의 다른 융합 단백질의 홉합을 포합하였고, 각 융합 단백질은 DGD, M12, PQAB 및 RSAB로 구성된 그룹으로부터 선별된 서로 다른 항원을 포함한다(도 13a). 마우스의 백신화 및 비장세포의 자극은 상기에 기술된 것과 비슷한 방법을 사용하여 수행하였다. 간단하게, 모든 마우스는 마지막 면역화 7일 후에 희생하였고, 비장세포를 수확하였다. 상기 비장세포를 분리하고, 1 ㎍/ml의 GolgiPlug(BD Pharmingen, San Diego, CA)의 존재하에서 비장세포를 자극할 수 있는 10 ㎍/ml의 개별적인 재조합 DGD, M12, PQAB 또는 RSAB 항원과 함께 6 웰 플레이트(2×107 세포/2 ml/웰)에서 37℃, 16시간 동안 배양하였다.
"DGD"(서열번호 26)의 아미노산 서열은 하기이다: RHHFTPSERQLCLSSIQTAFNQGAGTCILSDSGRISYTVEFSLPTHHTVRLIRVTAPPS ALDQVIRNALASPGSGGDLGEAIREQPEQARLALTLAAAESERFVRQGTGNDEAGAANADVVSLTCPVAAGECAGPADSGDALLERNYPTGAEFLGDGGDV RHHFTPSERQLCLSSIQTAFNQGAGTCILSDSGRISYTVEFSLPTHHTVRLIRVTAPPSA. DGD는 PRRSV ORF7 a.a.64 내지 a.a.123(굵은 글씨), 링커(밑줄친) 및 ORF7 a.a.64 내지 a.a.123(굻은 글씨)의 융합 항원을 나타낸다.
용어 "M12"는 PRRSV ORF1b a.a.1046 내지 a.a.1210의 항원을 나타낸다. 이것의 아미노산 서열(서열번호 27)은 하기와 같다: NNKECTVAQALGNGDKFRATDKRVVDSLRAICADLEGSSSPLPKVAHNLGFYFSPDLTQFAKLPIELAPHWPVVSTQNNEKWPDRLVASLRPLDKYSRACIGAGYMVGPSVFLGTPGVVSYYLTKFVKGEAQVLPETVFSTGRIEVDCREYLDDREREVAASLPH.
"PQAB"(서열번호 28)의 아미노산 서열은 하기와 같다: GSSLDDFCYDSTAPQKVLLAFSITYASNDSSSHLQLIYNLTLCELNGTDWLANKFDWA. PQAB는 PRRSV 미국 종 ORF6 a.a.2 내지 a.a.26 및 ORF5 a.a.31 내지 a.a.63(밑줄 친)의 융합 항원을 나타낸다.
RSAB의 아미노산 서열은 MGSLDDFCNDSTAAQKLVLAFSITYTPIFVAGGSSSTYQYIYNLTICELNGTDWLSNHFDWA(서열번호 29)이다. 용어 "RSAB"는 PRRSV 유럽 종 ORF6 a.a.2 내지 28 및 ORF5 a.a.31 내지 64(밑줄 친)의 융합 항원을 나타낸다.
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실험예
6>
융합 단백질 RAP1-CD28convPEt-PRRSV-K3의 RAP1 도메인 3의 절편은 융합 단백질 A2M-CD28convPEt-PRRSV-K3, Tat-CD28convPEt-PRRSV-K3 및 HSP-CD28convPEt-PRRSV-K3 백신을 생성하기 위해 각각 A2M 최소한(서열번호 6), HIV-Tat 최소한(서열번호 7) 또는 HSPs 최소한(서열번호 8)로 대체되었다. TC-1 종양 억제 활성 및 상기 백신으로 강화된 세포 매개 면역 반응은 상기에 기술한 비슷한 방법을 사용하여 검사하였다. 표 1은 다양한 융합 단백질의 요소의 서열번호를 나타낸다. 표 2는 동물에서 T 세포-매개의 면역 반응에 대한 효과에 대해 시험한 융합 단백질 및 항원의 서열을 나타낸다.
요소 | 서열번호 | 길이(잔기) |
hCD28 중심 TDIYFCKIEVMYPPPYLDNEKSNGTIIH |
1 |
28 |
hCD28 최대한 NCDGKLGNESVTFYLQNLYVNQTDIYFCKIEVMYPPPYLDNEKSNGTIIHVKG |
2 |
53 |
℃PEt 중심 (PE 전좌 도메인 중심; PE의 a.a.280 내지 a.a.313) |
3 |
34 |
PEt 최대한 (전좌 도메인 최대, PE의 a.a.253 내지 a.a.364) |
4 |
112 |
RAP1 최소한(RAP1의 도메인 III) |
5 |
104 |
A2M 최소한 |
6 |
153 |
HIV-Tat 최소한 |
7 |
24 |
HSPs 최소한, 열 충격(heat shock) 70 kDa 단백질(HSPs; 호모 사피언스) |
8 |
641 |
최소의 수도모나스 엑소톡신 A(Pseudomonas exotoxin A; PE) 결합 도메인 Ia(APC-결합 도메인, PE의 a.a.1 내지 a.a.252) |
9 |
252 |
링커 RXRXKR, 여기서 "X"는 임의의 아미노산 잔기 |
15 |
6 |
전장 PE(덱소톡신 A 성숙한(mature) 형태, 수도모나스 애루지노사(Pseudomonas aeruginosa)) |
10 |
613 |
전장 RAP1(호모 사피엔스 단백질 1과 연관된 낮은 밀도 지단백(lipoprotein) 수용체-연결된 단백질, LRPAP1); 도메인 1: a.a.1 내지 a.a.112; 도메인 2: a.a.113 내지 a.a.218; 도메인 3: a.a.219 내지 a.a.323. |
11 |
323 |
전장 A2M(호모 사피언스 알파-2-마크로글로불린(macroglobulin) 수용체-연관된 단백질 전구체(precursor)) |
12 |
357 |
HIV-Tat(인간 면역결핍 바이러스 1) |
13 |
101 |
KDEL |
14 |
4 |
KKDLRDELKDEL |
16 |
12 |
KKDELRDELKDEL |
17 |
13 |
KKDELRDELKDEL |
18 |
13 |
KKDELRDELKDEL |
19 |
12 |
PE268 -313 PLETFTRHRQPRGWEQLEQCGYPVQRLVALYLAARLSWNQVDQVIR |
20 |
46 |
CD28convPEt T1D2I3Y4F5C6K7X8E9X10X11Y12P13P14P15Y16X17D18N19E20K21S22N23G24T25I26I27H28R29X30R31X32K33R34G35W36E37Q38L39E40Q41C42G43Y44P45V46Q47R48L49V50A51L52Y53L54A55A56R57L58S59W60N61Q62V63D64Q65V66I67R68 여기서, (X)8은I 또는 L; (X)10은 V, F 또는 A, (X)11은 M 또는 L, X17는 L 또는 I, (X)30, 32는 임의의 아미노산 잔기 |
30 |
68 |
CD28 일치 서열 T1D2I3Y4F5C6K7(X)8E9(X)10(X)11Y12P13P14P15Y16X17D18N18E20K21S22N23G24T25I26I27H28 여기서, (X)8는I 또는 L; (X)10는 V, F 또는 A, (X)11는 M 또는 L, X17는 L 또는 I. |
31 |
28 |
CD28 주요 영역 K1(X)2E3(X)4(X)5Y6P7P8P9Y10, 여기서 (X)2는 I 또는 L; (X)4는 V, F 또는 A, (X)5는 M 또는 L. |
32 |
10 |
융합 단백질 명 | 항원 명 | 항원 서열번호 |
RAP1-CD28convPEt-E7-K3 |
HPV16 E7(전장) |
21 |
RAP1-CD28convPEt-E718-K3 |
HPV16 E7(전장) |
22 |
RAP1-CD28convPEt-HCVc-K3 |
HCV 중신 단백질(전장) |
23 |
RAP1-CD28convPEt-HBx-K3 |
HBV X 단백질(전장) |
24 |
RAP1-CD28convPEt-PCV2-K3 |
PCV2 ORF2(ORF2의 절편) |
25 |
RAP1-CD28convPEt-DGD-K3 |
PRRSV 뉴클레오갭시드(nucleocapsid) (융합 항원: ORF7 a.a.64 내지 a.a.123, 링커 및 ORF7 a.a.64 내지 a.a.123) |
26 |
RAP1-CD28convPEt-M12-K3 |
PRRSV RNA-의존 RNA 중합효소(polymerase) (ORF1b a.a.1046 내지 a.a.1210) |
27 |
RAP1-CD28convPEt-PQAB-K3 |
PRRSV 미국 종: ORF6(a.a.2 내지 a.a.26) 및 ORF5(a.a.31 내지 a.a.63)의 융합 항원 |
28 |
RAP1-CD28convPEt-RSAB-K3 |
PRRSV 유럽 종: ORF6(a.a.2 내지 a.a.28) 및 ORF5(a.a.31 내지 a.a.64)의 융합 항원 |
29 |
면역원성 분석에서, 다양한 백신에 의해 유도된 항원-특이적인 세포-매개의 면역 반응은 비장세포에서 CD3+/CD4+/IFNγ+ 및 CD3+/CD8+/IFNγ+ T 세포의 개수를 측정하여 평가하였다. 상기 결과는 백신 RAP1-CD28convPEt-항원-K3가 강한 T 세포 반응을 유도할 수 있음을 나타낸다. 도 14b는 CD28convPEt-E718-K3에 유도된 CD3+/CD4+/IFNγ+ T 세포의 개수 및 CD3+/CD8+/IFNγ+ T 세포의 개수가 PE407-E716-K3에 비해 각각 5배 및 7배임을 나타낸다. 이는 백신 RAP1-CD28convPEt-E7-K3가 PE407-E7-K3에 비해 더 나은 세포-매개의 면역원성을 갖는다는 것을 나타낸다.
RAP1 도메인 III, 전좌 펩타이드 PEt가 없는 민감성 신호 CD28conv 단독, 항원 및 ER 유지 신호를 포함하는 융합 단백질은 항원이 열 개 또는 10개 이상의 항원결정기를 포함하도록 선택되었을 때 강한 항원-특정 T 세포 매개의 면역 반응을 유도하는 데에 충분하다. 도 14c는 백신 RAP1-CD28conv-HCVcore-K3가 플라시보 그룹에 비해 각각 20배 및 7.6배의 CD3+/CD4+/IFNγ+ 및 CD3+/CD8+/IFNγ+ T 세포의 개수를 갖는 T 세포 반응을 유도함을 나타낸다. 항원 HCVcore은 11개의 잘 알려진 MHC I 항원결정기를 포함한다.
강한 세포-매개의 면역원성을 유도하기 위해 본 발명의 융합 단백질에 대해서 ER 유지 신호가 필수적이지 않다는 것은 예상 외의 결과였다. 다른 말로, ER 유지 서열 없이, 본 발명의 융합 단백질이 강한 T-세포 반응을 유도할 수 있다. 도 14d는 RAP1-CD28convPEt-HBx(ER 유지 신호 K3 없는)에 의해 유도된 CD3+/CD4+/IFNγ+ 및 CD3+/CD8+/IFNγ+ T 세포의 개수가 플라시보에 비해서 7배 및 74배임을 나타낸다.
반면, 미국 특허 번호 7378100B2 및 7335361은 ER 유지 신호 K3가 T 세포 반응을 유도하기 위해서 PE-연관된 융합 단백질(PE407-항원-K3)에 대해 필수적임을 보여준다.
또한 RAP1 도메인 III, 전좌 펩타이드 PE218 -313(민감성 신호 CD28conv가 없는), 항원 및 ER 유지 신호를 포함하는 융합 단백질이 RAP1 도메인 III를 포함하지 않는 PE-연관된 융합 단백질보다 우수하다는 것을 발견하였다. 도 15c 내지 j는 백신 RAP1-PE268 -313-PRRSV-K3가 PE407-PRRSV-K3보다 더 큰 CD3+/CD4+/IFNγ+ 및 CD3+/CD8+/IFNγ+ T 세포의 수를 유도함을 나타내었다.
본 발명의 예시적인 실시예의 앞선 기술은 오직 설명 및 기술의 목적을 위해서 제시되며 개시된 자세한 형태로 본 발명에 대해 철저하게 또는 한정하기 위해 의도되지 않는다. 많은 변형 및 변화가 상기 교시(teaching)의 관점에서 가능하다. 상기 실시예 및 실험예는 고려되는 특정한 사용에 적합한 바와 같이 당업계의 숙련된 다른 사람들이 본 발명 및 다양한 변형과 함께 다양한 실시예를 활용하도록 본 발명의 원리 및 그 실제 적용을 설명하기 위하여 선택되고 기술되었다. 대안적인 실시예는 본 발명의 정신 및 범위로부터 벗어나는 것 없이, 발명이 속하는 기술분야에서 통상의 지식을 가진 자에게 명백해 질 것이다. 따라서, 본 발명의 범위는 앞선 기술 및 본 명세서에 기술된 예시적인 실시예 보다는 첨부된 청구항에 의해 정의된다.
<110> TheVax Genetics Vaccine Co., Ltd
WU, CHIA-MAO
CHANG, HSIU-KANG
<120> FUSION PROTEINS AS IMMUNOGEN ENHANCERS FOR ENHANCING
ANTIGEN-SPECIFIC T CELL RESPONSES
<130> 10021-00001
<150> US/61733879
<151> 2012-12-05
<160> 32
<170> PatentIn version 3.5
<210> 1
<211> 28
<212> PRT
<213> Artificial Sequence
<220>
<223> hCD28 Core
<400> 1
Thr Asp Ile Tyr Phe Cys Lys Ile Glu Val Met Tyr Pro Pro Pro Tyr
1 5 10 15
Leu Asp Asn Glu Lys Ser Asn Gly Thr Ile Ile His
20 25
<210> 2
<211> 53
<212> PRT
<213> Artificial Sequence
<220>
<223> hCD28 Maximum
<400> 2
Asn Cys Asp Gly Lys Leu Gly Asn Glu Ser Val Thr Phe Tyr Leu Gln
1 5 10 15
Asn Leu Tyr Val Asn Gln Thr Asp Ile Tyr Phe Cys Lys Ile Glu Val
20 25 30
Met Tyr Pro Pro Pro Tyr Leu Asp Asn Glu Lys Ser Asn Gly Thr Ile
35 40 45
Ile His Val Lys Gly
50
<210> 3
<211> 34
<212> PRT
<213> Artificial Sequence
<220>
<223> PEt Core
<400> 3
Gly Trp Glu Gln Leu Glu Gln Cys Gly Tyr Pro Val Gln Arg Leu Val
1 5 10 15
Ala Leu Tyr Leu Ala Ala Arg Leu Ser Trp Asn Gln Val Asp Gln Val
20 25 30
Ile Arg
<210> 4
<211> 112
<212> PRT
<213> Artificial Sequence
<220>
<223> PEt Maximum
<400> 4
Gly Gly Ser Leu Ala Ala Leu Thr Ala His Gln Ala Cys His Leu Pro
1 5 10 15
Leu Glu Thr Phe Thr Arg His Arg Gln Pro Arg Gly Trp Glu Gln Leu
20 25 30
Glu Gln Cys Gly Tyr Pro Val Gln Arg Leu Val Ala Leu Tyr Leu Ala
35 40 45
Ala Arg Leu Ser Trp Asn Gln Val Asp Gln Val Ile Arg Asn Ala Leu
50 55 60
Ala Ser Pro Gly Ser Gly Gly Asp Leu Gly Glu Ala Ile Arg Glu Gln
65 70 75 80
Pro Glu Gln Ala Arg Leu Ala Leu Thr Leu Ala Ala Ala Glu Ser Glu
85 90 95
Arg Phe Val Arg Gln Gly Thr Gly Asn Asp Glu Ala Gly Ala Ala Asn
100 105 110
<210> 5
<211> 104
<212> PRT
<213> Artificial Sequence
<220>
<223> RAP1 Minimum
<400> 5
Ala Glu Phe Glu Glu Pro Arg Val Ile Asp Leu Trp Asp Leu Ala Gln
1 5 10 15
Ser Ala Asn Leu Thr Asp Lys Glu Leu Glu Ala Phe Arg Glu Glu Leu
20 25 30
Lys His Phe Glu Ala Lys Ile Glu Lys His Asn His Tyr Gln Lys Gln
35 40 45
Leu Glu Ile Ala His Glu Lys Leu Arg His Ala Glu Ser Val Gly Asp
50 55 60
Gly Glu Arg Val Ser Arg Ser Arg Glu Lys His Ala Leu Leu Glu Gly
65 70 75 80
Arg Thr Lys Glu Leu Gly Tyr Thr Val Lys Lys His Leu Gln Asp Leu
85 90 95
Ser Gly Arg Ile Ser Arg Ala Arg
100
<210> 6
<211> 153
<212> PRT
<213> Artificial Sequence
<220>
<223> A2M Minimum
<400> 6
Val Tyr Leu Gln Thr Ser Leu Lys Tyr Asn Ile Leu Pro Glu Lys Glu
1 5 10 15
Glu Phe Pro Phe Ala Leu Gly Val Gln Thr Leu Pro Gln Thr Cys Asp
20 25 30
Glu Pro Lys Ala His Thr Ser Phe Gln Ile Ser Leu Ser Val Ser Tyr
35 40 45
Thr Gly Ser Arg Ser Ala Ser Asn Met Ala Ile Val Asp Val Lys Met
50 55 60
Val Ser Gly Phe Ile Pro Leu Lys Pro Thr Val Lys Met Leu Glu Arg
65 70 75 80
Ser Asn His Val Ser Arg Thr Glu Val Ser Ser Asn His Val Leu Ile
85 90 95
Tyr Leu Asp Lys Val Ser Asn Gln Thr Leu Ser Leu Phe Phe Thr Val
100 105 110
Leu Gln Asp Val Pro Val Arg Asp Leu Lys Pro Ala Ile Val Lys Val
115 120 125
Tyr Asp Tyr Tyr Glu Thr Asp Glu Phe Ala Ile Ala Glu Tyr Asn Ala
130 135 140
Pro Cys Ser Lys Asp Leu Gly Asn Ala
145 150
<210> 7
<211> 24
<212> PRT
<213> Artificial Sequence
<220>
<223> HIV-Tat Minimum
<400> 7
Arg Gly Asp Pro Thr Gly Gln Glu Glu Ser Lys Glu Lys Val Glu Lys
1 5 10 15
Glu Thr Val Val Asp Pro Val Thr
20
<210> 8
<211> 641
<212> PRT
<213> Artificial Sequence
<220>
<223> HSPs Minimum
<400> 8
Met Ala Lys Ala Ala Ala Ile Gly Ile Asp Leu Gly Thr Thr Tyr Ser
1 5 10 15
Cys Val Gly Val Phe Gln His Gly Lys Val Glu Ile Ile Ala Asn Asp
20 25 30
Gln Gly Asn Arg Thr Thr Pro Ser Tyr Val Ala Phe Thr Asp Thr Glu
35 40 45
Arg Leu Ile Gly Asp Ala Ala Lys Asn Gln Val Ala Leu Asn Pro Gln
50 55 60
Asn Thr Val Phe Asp Ala Lys Arg Leu Ile Gly Arg Lys Phe Gly Asp
65 70 75 80
Pro Val Val Gln Ser Asp Met Lys His Trp Pro Phe Gln Val Ile Asn
85 90 95
Asp Gly Asp Lys Pro Lys Val Gln Val Ser Tyr Lys Gly Glu Thr Lys
100 105 110
Ala Phe Tyr Pro Glu Glu Ile Ser Ser Met Val Leu Thr Lys Met Lys
115 120 125
Glu Ile Ala Glu Ala Tyr Leu Gly Tyr Pro Val Thr Asn Ala Val Ile
130 135 140
Thr Val Pro Ala Tyr Phe Asn Asp Ser Gln Arg Gln Ala Thr Lys Asp
145 150 155 160
Ala Gly Val Ile Ala Gly Leu Asn Val Leu Arg Ile Ile Asn Glu Pro
165 170 175
Thr Ala Ala Ala Ile Ala Tyr Gly Leu Asp Arg Thr Gly Lys Gly Glu
180 185 190
Arg Asn Val Leu Ile Phe Asp Leu Gly Gly Gly Thr Phe Asp Val Ser
195 200 205
Ile Leu Thr Ile Asp Asp Gly Ile Phe Glu Val Lys Ala Thr Ala Gly
210 215 220
Asp Thr His Leu Gly Gly Glu Asp Phe Asp Asn Arg Leu Val Asn His
225 230 235 240
Phe Val Glu Glu Phe Lys Arg Lys His Lys Lys Asp Ile Ser Gln Asn
245 250 255
Lys Arg Ala Val Arg Arg Leu Arg Thr Ala Cys Glu Arg Ala Lys Arg
260 265 270
Thr Leu Ser Ser Ser Thr Gln Ala Ser Leu Glu Ile Asp Ser Leu Phe
275 280 285
Glu Gly Ile Asp Phe Tyr Thr Ser Ile Thr Arg Ala Arg Phe Glu Glu
290 295 300
Leu Cys Ser Asp Leu Phe Arg Ser Thr Leu Glu Pro Val Glu Lys Ala
305 310 315 320
Leu Arg Asp Ala Lys Leu Asp Lys Ala Gln Ile His Asp Leu Val Leu
325 330 335
Val Gly Gly Ser Thr Arg Ile Pro Lys Val Gln Lys Leu Leu Gln Asp
340 345 350
Phe Phe Asn Gly Arg Asp Leu Asn Lys Ser Ile Asn Pro Asp Glu Ala
355 360 365
Val Ala Tyr Gly Ala Ala Val Gln Ala Ala Ile Leu Met Gly Asp Lys
370 375 380
Ser Glu Asn Val Gln Asp Leu Leu Leu Leu Asp Val Ala Pro Leu Ser
385 390 395 400
Leu Gly Leu Glu Thr Ala Gly Gly Val Met Thr Ala Leu Ile Lys Arg
405 410 415
Asn Ser Thr Ile Pro Thr Lys Gln Thr Gln Ile Phe Thr Thr Tyr Ser
420 425 430
Asp Asn Gln Pro Gly Val Leu Ile Gln Val Tyr Glu Gly Glu Arg Ala
435 440 445
Met Thr Lys Asp Asn Asn Leu Leu Gly Arg Phe Glu Leu Ser Gly Ile
450 455 460
Pro Pro Ala Pro Arg Gly Val Pro Gln Ile Glu Val Thr Phe Asp Ile
465 470 475 480
Asp Ala Asn Gly Ile Leu Asn Val Thr Ala Thr Asp Lys Ser Thr Gly
485 490 495
Lys Ala Asn Lys Ile Thr Ile Thr Asn Asp Lys Gly Arg Leu Ser Lys
500 505 510
Glu Glu Ile Glu Arg Met Val Gln Glu Ala Glu Lys Tyr Lys Ala Glu
515 520 525
Asp Glu Val Gln Arg Glu Arg Val Ser Ala Lys Asn Ala Leu Glu Ser
530 535 540
Tyr Ala Phe Asn Met Lys Ser Ala Val Glu Asp Glu Gly Leu Lys Gly
545 550 555 560
Lys Ile Ser Glu Ala Asp Lys Lys Lys Val Leu Asp Lys Cys Gln Glu
565 570 575
Val Ile Ser Trp Leu Asp Ala Asn Thr Leu Ala Glu Lys Asp Glu Phe
580 585 590
Glu His Lys Arg Lys Glu Leu Glu Gln Val Cys Asn Pro Ile Ile Ser
595 600 605
Gly Leu Tyr Gln Gly Ala Gly Gly Pro Gly Pro Gly Gly Phe Gly Ala
610 615 620
Gln Gly Pro Lys Gly Gly Ser Gly Ser Gly Pro Thr Ile Glu Glu Val
625 630 635 640
Asp
<210> 9
<211> 252
<212> PRT
<213> Pseudomonas aeruginosa
<400> 9
Ala Glu Glu Ala Phe Asp Leu Trp Asn Glu Cys Ala Lys Ala Cys Val
1 5 10 15
Leu Asp Leu Lys Asp Gly Val Arg Ser Ser Arg Met Ser Val Asp Pro
20 25 30
Ala Ile Ala Asp Thr Asn Gly Gln Gly Val Leu His Tyr Ser Met Val
35 40 45
Leu Glu Gly Gly Asn Asp Ala Leu Lys Leu Ala Ile Asp Asn Ala Leu
50 55 60
Ser Ile Thr Ser Asp Gly Leu Thr Ile Arg Leu Glu Gly Gly Val Glu
65 70 75 80
Pro Asn Lys Pro Val Arg Tyr Ser Tyr Thr Arg Gln Ala Arg Gly Ser
85 90 95
Trp Ser Leu Asn Trp Leu Val Pro Ile Gly His Glu Lys Pro Ser Asn
100 105 110
Ile Lys Val Phe Ile His Glu Leu Asn Ala Gly Asn Gln Leu Ser His
115 120 125
Met Ser Pro Ile Tyr Thr Ile Glu Met Gly Asp Glu Leu Leu Ala Lys
130 135 140
Leu Ala Arg Asp Ala Thr Phe Phe Val Arg Ala His Glu Ser Asn Glu
145 150 155 160
Met Gln Pro Thr Leu Ala Ile Ser His Ala Gly Val Ser Val Val Met
165 170 175
Ala Gln Thr Gln Pro Arg Arg Glu Lys Arg Trp Ser Glu Trp Ala Ser
180 185 190
Gly Lys Val Leu Cys Leu Leu Asp Pro Leu Asp Gly Val Tyr Asn Tyr
195 200 205
Leu Ala Gln Gln Arg Cys Asn Leu Asp Asp Thr Trp Glu Gly Lys Ile
210 215 220
Tyr Arg Val Leu Ala Gly Asn Pro Ala Lys His Asp Leu Asp Ile Lys
225 230 235 240
Pro Thr Val Ile Ser His Arg Leu His Phe Pro Glu
245 250
<210> 10
<211> 613
<212> PRT
<213> Pseudomonas aeruginosa
<400> 10
Ala Glu Glu Ala Phe Asp Leu Trp Asn Glu Cys Ala Lys Ala Cys Val
1 5 10 15
Leu Asp Leu Lys Asp Gly Val Arg Ser Ser Arg Met Ser Val Asp Pro
20 25 30
Ala Ile Ala Asp Thr Asn Gly Gln Gly Val Leu His Tyr Ser Met Val
35 40 45
Leu Glu Gly Gly Asn Asp Ala Leu Lys Leu Ala Ile Asp Asn Ala Leu
50 55 60
Ser Ile Thr Ser Asp Gly Leu Thr Ile Arg Leu Glu Gly Gly Val Glu
65 70 75 80
Pro Asn Lys Pro Val Arg Tyr Ser Tyr Thr Arg Gln Ala Arg Gly Ser
85 90 95
Trp Ser Leu Asn Trp Leu Val Pro Ile Gly His Glu Lys Pro Ser Asn
100 105 110
Ile Lys Val Phe Ile His Glu Leu Asn Ala Gly Asn Gln Leu Ser His
115 120 125
Met Ser Pro Ile Tyr Thr Ile Glu Met Gly Asp Glu Leu Leu Ala Lys
130 135 140
Leu Ala Arg Asp Ala Thr Phe Phe Val Arg Ala His Glu Ser Asn Glu
145 150 155 160
Met Gln Pro Thr Leu Ala Ile Ser His Ala Gly Val Ser Val Val Met
165 170 175
Ala Gln Thr Gln Pro Arg Arg Glu Lys Arg Trp Ser Glu Trp Ala Ser
180 185 190
Gly Lys Val Leu Cys Leu Leu Asp Pro Leu Asp Gly Val Tyr Asn Tyr
195 200 205
Leu Ala Gln Gln Arg Cys Asn Leu Asp Asp Thr Trp Glu Gly Lys Ile
210 215 220
Tyr Arg Val Leu Ala Gly Asn Pro Ala Lys His Asp Leu Asp Ile Lys
225 230 235 240
Pro Thr Val Ile Ser His Arg Leu His Phe Pro Glu Gly Gly Ser Leu
245 250 255
Ala Ala Leu Thr Ala His Gln Ala Cys His Leu Pro Leu Glu Thr Phe
260 265 270
Thr Arg His Arg Gln Pro Arg Gly Trp Glu Gln Leu Glu Gln Cys Gly
275 280 285
Tyr Pro Val Gln Arg Leu Val Ala Leu Tyr Leu Ala Ala Arg Leu Ser
290 295 300
Trp Asn Gln Val Asp Gln Val Ile Arg Asn Ala Leu Ala Ser Pro Gly
305 310 315 320
Ser Gly Gly Asp Leu Gly Glu Ala Ile Arg Glu Gln Pro Glu Gln Ala
325 330 335
Arg Leu Ala Leu Thr Leu Ala Ala Ala Glu Ser Glu Arg Phe Val Arg
340 345 350
Gln Gly Thr Gly Asn Asp Glu Ala Gly Ala Ala Asn Ala Asp Val Val
355 360 365
Ser Leu Thr Cys Pro Val Ala Ala Gly Glu Cys Ala Gly Pro Ala Asp
370 375 380
Ser Gly Asp Ala Leu Leu Glu Arg Asn Tyr Pro Thr Gly Ala Glu Phe
385 390 395 400
Leu Gly Asp Gly Gly Asp Val Ser Phe Ser Thr Arg Gly Thr Gln Asn
405 410 415
Trp Thr Val Glu Arg Leu Leu Gln Ala His Arg Gln Leu Glu Glu Arg
420 425 430
Gly Tyr Val Phe Val Gly Tyr His Gly Thr Phe Leu Glu Ala Ala Gln
435 440 445
Ser Ile Val Phe Gly Gly Val Arg Ala Arg Ser Gln Asp Leu Asp Ala
450 455 460
Ile Trp Arg Gly Phe Tyr Ile Ala Gly Asp Pro Ala Leu Ala Tyr Gly
465 470 475 480
Tyr Ala Gln Asp Gln Glu Pro Asp Ala Arg Gly Arg Ile Arg Asn Gly
485 490 495
Ala Leu Leu Arg Val Tyr Val Pro Arg Ser Ser Leu Pro Gly Phe Tyr
500 505 510
Arg Thr Ser Leu Thr Leu Ala Ala Pro Glu Ala Ala Gly Glu Val Glu
515 520 525
Arg Leu Ile Gly His Pro Leu Pro Leu Arg Leu Asp Ala Ile Thr Gly
530 535 540
Pro Glu Glu Glu Gly Gly Arg Leu Glu Thr Ile Leu Gly Trp Pro Leu
545 550 555 560
Ala Glu Arg Thr Val Val Ile Pro Ser Ala Ile Pro Thr Asp Pro Arg
565 570 575
Asn Val Gly Gly Asp Leu Asp Pro Ser Ser Ile Pro Asp Lys Glu Gln
580 585 590
Ala Ile Ser Ala Leu Pro Asp Tyr Ala Ser Gln Pro Gly Lys Pro Pro
595 600 605
Arg Glu Asp Leu Lys
610
<210> 11
<211> 323
<212> PRT
<213> Homo sapiens
<400> 11
Tyr Ser Arg Glu Lys Asn Gln Pro Lys Pro Ser Pro Lys Arg Glu Ser
1 5 10 15
Gly Glu Glu Phe Arg Met Glu Lys Leu Asn Gln Leu Trp Glu Lys Ala
20 25 30
Gln Arg Leu His Leu Pro Pro Val Arg Leu Ala Glu Leu His Ala Asp
35 40 45
Leu Lys Ile Gln Glu Arg Asp Glu Leu Ala Trp Lys Lys Leu Lys Leu
50 55 60
Asp Gly Leu Asp Glu Asp Gly Glu Lys Glu Ala Arg Leu Ile Arg Asn
65 70 75 80
Leu Asn Val Ile Leu Ala Lys Tyr Gly Leu Asp Gly Lys Lys Asp Ala
85 90 95
Arg Gln Val Thr Ser Asn Ser Leu Ser Gly Thr Gln Glu Asp Gly Leu
100 105 110
Asp Asp Pro Arg Leu Glu Lys Leu Trp His Lys Ala Lys Thr Ser Gly
115 120 125
Lys Phe Ser Gly Glu Glu Leu Asp Lys Leu Trp Arg Glu Phe Leu His
130 135 140
His Lys Glu Lys Val His Glu Tyr Asn Val Leu Leu Glu Thr Leu Ser
145 150 155 160
Arg Thr Glu Glu Ile His Glu Asn Val Ile Ser Pro Ser Asp Leu Ser
165 170 175
Asp Ile Lys Gly Ser Val Leu His Ser Arg His Thr Glu Leu Lys Glu
180 185 190
Lys Leu Arg Ser Ile Asn Gln Gly Leu Asp Arg Leu Arg Arg Val Ser
195 200 205
His Gln Gly Tyr Ser Thr Glu Ala Glu Phe Glu Glu Pro Arg Val Ile
210 215 220
Asp Leu Trp Asp Leu Ala Gln Ser Ala Asn Leu Thr Asp Lys Glu Leu
225 230 235 240
Glu Ala Phe Arg Glu Glu Leu Lys His Phe Glu Ala Lys Ile Glu Lys
245 250 255
His Asn His Tyr Gln Lys Gln Leu Glu Ile Ala His Glu Lys Leu Arg
260 265 270
His Ala Glu Ser Val Gly Asp Gly Glu Arg Val Ser Arg Ser Arg Glu
275 280 285
Lys His Ala Leu Leu Glu Gly Arg Thr Lys Glu Leu Gly Tyr Thr Val
290 295 300
Lys Lys His Leu Gln Asp Leu Ser Gly Arg Ile Ser Arg Ala Arg His
305 310 315 320
Asn Glu Leu
<210> 12
<211> 357
<212> PRT
<213> Homo sapiens
<400> 12
Met Ala Pro Arg Arg Val Arg Ser Phe Leu Arg Gly Leu Pro Ala Leu
1 5 10 15
Leu Leu Leu Leu Leu Phe Leu Gly Pro Trp Pro Ala Ala Ser His Gly
20 25 30
Gly Lys Tyr Ser Arg Glu Lys Asn Gln Pro Lys Pro Ser Pro Lys Arg
35 40 45
Glu Ser Gly Glu Glu Phe Arg Met Glu Lys Leu Asn Gln Leu Trp Glu
50 55 60
Lys Ala Gln Arg Leu His Leu Pro Pro Val Arg Leu Ala Glu Leu His
65 70 75 80
Ala Asp Leu Lys Ile Gln Glu Arg Asp Glu Leu Ala Trp Lys Lys Leu
85 90 95
Lys Leu Asp Gly Leu Asp Glu Asp Gly Glu Lys Glu Ala Arg Leu Ile
100 105 110
Arg Asn Leu Asn Val Ile Leu Ala Lys Tyr Gly Leu Asp Gly Lys Lys
115 120 125
Asp Ala Arg Gln Val Thr Ser Asn Ser Leu Ser Gly Thr Gln Glu Asp
130 135 140
Gly Leu Asp Asp Pro Arg Leu Glu Lys Leu Trp His Lys Ala Lys Thr
145 150 155 160
Ser Gly Lys Phe Ser Gly Glu Glu Leu Asp Lys Leu Trp Arg Glu Phe
165 170 175
Leu His His Lys Glu Lys Val His Glu Tyr Asn Val Leu Leu Glu Thr
180 185 190
Leu Ser Arg Thr Glu Glu Ile His Glu Asn Val Ile Ser Pro Ser Asp
195 200 205
Leu Ser Asp Ile Lys Gly Ser Val Leu His Ser Arg His Thr Glu Leu
210 215 220
Lys Glu Lys Leu Arg Ser Ile Asn Gln Gly Leu Asp Arg Leu Arg Arg
225 230 235 240
Val Ser His Gln Gly Tyr Ser Thr Glu Ala Glu Phe Glu Glu Pro Arg
245 250 255
Val Ile Asp Leu Trp Asp Leu Ala Gln Ser Ala Asn Leu Thr Asp Lys
260 265 270
Glu Leu Glu Ala Phe Arg Glu Glu Leu Lys His Phe Glu Ala Lys Ile
275 280 285
Glu Lys His Asn His Tyr Gln Lys Gln Leu Glu Ile Ala His Glu Lys
290 295 300
Leu Arg His Ala Glu Ser Val Gly Asp Gly Glu Arg Val Ser Arg Ser
305 310 315 320
Arg Glu Lys His Ala Leu Leu Glu Gly Arg Thr Lys Glu Leu Gly Tyr
325 330 335
Thr Val Lys Lys His Leu Gln Asp Leu Ser Gly Arg Ile Ser Arg Ala
340 345 350
Arg His Asn Glu Leu
355
<210> 13
<211> 101
<212> PRT
<213> Human immunodeficiency virus
<400> 13
Met Glu Pro Val Asp Pro Arg Leu Glu Pro Trp Lys His Pro Gly Ser
1 5 10 15
Gln Pro Lys Thr Pro Cys Thr Lys Cys Tyr Cys Lys Lys Cys Cys Leu
20 25 30
His Cys Gln Val Cys Phe Met Thr Lys Gly Leu Gly Ile Ser Tyr Gly
35 40 45
Arg Lys Lys Arg Arg Gln Arg Arg Arg Ala Pro Gln Asp Asn Lys Asn
50 55 60
His Gln Val Ser Leu Ser Lys Gln Pro Thr Ser Arg Ala Arg Gly Asp
65 70 75 80
Pro Thr Gly Gln Glu Glu Ser Lys Glu Lys Val Glu Lys Glu Thr Val
85 90 95
Val Asp Pro Val Thr
100
<210> 14
<211> 4
<212> PRT
<213> Artificial Sequence
<220>
<223> ER retention sequence
<400> 14
Lys Asp Glu Leu
1
<210> 15
<211> 6
<212> PRT
<213> Artificial Sequence
<220>
<223> Linker of CD28-PEt
<220>
<221> MISC_FEATURE
<222> (2)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (4)
<223> Xaa can be any naturally occurring amino acid
<400> 15
Arg Xaa Arg Xaa Lys Arg
1 5
<210> 16
<211> 12
<212> PRT
<213> Artificial Sequence
<220>
<223> ER retention sequence
<400> 16
Lys Lys Asp Leu Arg Asp Glu Leu Lys Asp Glu Leu
1 5 10
<210> 17
<211> 13
<212> PRT
<213> Artificial Sequence
<220>
<223> ER retention sequence
<400> 17
Lys Lys Asp Glu Leu Arg Asp Glu Leu Lys Asp Glu Leu
1 5 10
<210> 18
<211> 13
<212> PRT
<213> Artificial Sequence
<220>
<223> ER retention sequence
<400> 18
Lys Lys Asp Glu Leu Arg Val Glu Leu Lys Asp Glu Leu
1 5 10
<210> 19
<211> 12
<212> PRT
<213> Artificial Sequence
<220>
<223> ER retention sequence
<400> 19
Lys Asp Glu Leu Lys Asp Glu Leu Lys Asp Glu Leu
1 5 10
<210> 20
<211> 46
<212> PRT
<213> Pseudomonas aeruginosa
<400> 20
Pro Leu Glu Thr Phe Thr Arg His Arg Gln Pro Arg Gly Trp Glu Gln
1 5 10 15
Leu Glu Gln Cys Gly Tyr Pro Val Gln Arg Leu Val Ala Leu Tyr Leu
20 25 30
Ala Ala Arg Leu Ser Trp Asn Gln Val Asp Gln Val Ile Arg
35 40 45
<210> 21
<211> 98
<212> PRT
<213> Human papillomavirus type 16
<400> 21
Met His Gly Asp Thr Pro Thr Leu His Glu Tyr Met Leu Asp Leu Gln
1 5 10 15
Pro Glu Thr Thr Asp Leu Tyr Cys Tyr Glu Gln Leu Asn Asp Ser Ser
20 25 30
Glu Glu Glu Asp Glu Ile Asp Gly Pro Ala Gly Gln Ala Glu Pro Asp
35 40 45
Arg Ala His Tyr Asn Ile Val Thr Phe Cys Cys Lys Cys Asp Ser Thr
50 55 60
Leu Arg Leu Cys Val Gln Ser Thr His Val Asp Ile Arg Thr Leu Glu
65 70 75 80
Asp Leu Leu Met Gly Thr Leu Gly Ile Val Cys Pro Ile Cys Ser Gln
85 90 95
Lys Pro
<210> 22
<211> 105
<212> PRT
<213> Human papillomavirus type 18
<400> 22
Met His Gly Pro Lys Ala Thr Leu Gln Asp Ile Val Leu His Leu Glu
1 5 10 15
Pro Gln Asn Glu Ile Pro Val Asp Leu Leu Cys His Glu Gln Leu Ser
20 25 30
Asp Ser Glu Glu Glu Asn Asp Glu Ile Asp Gly Val Asn His Gln His
35 40 45
Leu Pro Ala Arg Arg Ala Glu Pro Gln Arg His Thr Met Leu Cys Met
50 55 60
Cys Cys Lys Cys Glu Ala Arg Ile Lys Leu Val Val Glu Ser Ser Ala
65 70 75 80
Asp Asp Leu Arg Ala Phe Gln Gln Leu Phe Leu Asn Thr Leu Ser Phe
85 90 95
Val Cys Pro Trp Cys Ala Ser Gln Gln
100 105
<210> 23
<211> 190
<212> PRT
<213> Hepatitis C virus
<400> 23
Met Ser Thr Asn Pro Lys Pro Gln Arg Lys Thr Lys Arg Asn Thr Asn
1 5 10 15
Arg Arg Pro Gln Asp Val Lys Phe Pro Gly Gly Gly Gln Ile Val Gly
20 25 30
Gly Val Tyr Leu Leu Pro Arg Arg Gly Pro Arg Leu Gly Val Arg Ala
35 40 45
Thr Arg Lys Thr Ser Glu Arg Ser Gln Pro Arg Gly Arg Arg Gln Pro
50 55 60
Ile Pro Lys Ala Arg Arg Pro Glu Gly Arg Thr Trp Ala Gln Pro Gly
65 70 75 80
Tyr Pro Trp Pro Leu Tyr Gly Asn Glu Gly Met Gly Trp Ala Gly Trp
85 90 95
Leu Leu Ser Pro Arg Gly Ser Arg Pro Asn Trp Gly Pro Thr Asp Pro
100 105 110
Arg Arg Arg Ser Arg Asn Leu Gly Lys Val Ile Asp Thr Leu Thr Cys
115 120 125
Gly Phe Ala Asp Leu Met Gly Tyr Ile Pro Leu Val Gly Ala Pro Leu
130 135 140
Gly Gly Val Ala Arg Ala Leu Ala His Gly Val Arg Val Leu Glu Asp
145 150 155 160
Gly Val Asn Tyr Ala Thr Gly Asn Leu Pro Gly Cys Ser Phe Ser Ile
165 170 175
Phe Leu Leu Ala Leu Leu Ser Cys Leu Thr Thr Pro Ala Ser
180 185 190
<210> 24
<211> 154
<212> PRT
<213> Hepatitis B virus
<400> 24
Met Ala Ala Arg Met Cys Cys Gln Leu Asp Pro Ala Arg Asp Val Leu
1 5 10 15
Cys Leu Arg Pro Val Gly Ala Glu Ser Arg Gly Arg Pro Leu Pro Gly
20 25 30
Pro Leu Gly Ala Leu Pro Pro Ser Ser Ala Ser Ala Val Pro Ala Asp
35 40 45
His Gly Ser His Leu Ser Leu Arg Gly Leu Pro Val Cys Ser Phe Ser
50 55 60
Ser Ala Gly Pro Cys Ala Leu Arg Phe Thr Ser Ala Arg Arg Met Glu
65 70 75 80
Thr Thr Val Asn Ala Pro Trp Ser Leu Pro Thr Val Leu His Lys Arg
85 90 95
Thr Ile Gly Leu Ser Gly Arg Ser Met Thr Trp Ile Glu Glu Tyr Ile
100 105 110
Lys Asp Cys Val Phe Lys Asp Trp Glu Glu Leu Gly Glu Glu Ile Arg
115 120 125
Leu Lys Val Phe Val Leu Gly Gly Cys Arg His Lys Leu Val Cys Ser
130 135 140
Pro Ala Pro Cys Asn Phe Phe Thr Ser Ala
145 150
<210> 25
<211> 192
<212> PRT
<213> Porcine circovirus
<400> 25
Asn Gly Ile Phe Asn Thr Arg Leu Ser Arg Thr Phe Gly Tyr Thr Ile
1 5 10 15
Lys Arg Thr Thr Val Lys Thr Pro Ser Trp Ala Val Asp Met Met Arg
20 25 30
Phe Asn Ile Asn Asp Phe Leu Pro Pro Gly Gly Gly Ser Asn Pro Arg
35 40 45
Ser Val Pro Phe Glu Tyr Tyr Ser Ile Ser Lys Val Lys Val Glu Phe
50 55 60
Trp Pro Cys Ser Pro Ile Thr Gln Gly Asp Ser Gly Val Gly Ser Ser
65 70 75 80
Ala Val Ile Leu Asp Asp Asn Phe Val Thr Lys Ala Thr Ala Leu Thr
85 90 95
Tyr Asp Pro Tyr Val Asn Tyr Ser Ser Arg His Thr Ile Thr Gln Pro
100 105 110
Phe Ser Tyr His Ser Arg Tyr Phe Thr Pro Lys Pro Val Leu Asp Ser
115 120 125
Thr Ile Asp Tyr Phe Gln Pro Asn Asn Lys Arg Asn Gln Leu Trp Leu
130 135 140
Arg Leu Gln Thr Ala Gly Asn Val Asp His Val Gly Leu Gly Thr Ala
145 150 155 160
Phe Glu Asn Ser Ile Tyr Asp Gln Glu Tyr Asn Ile Arg Val Thr Met
165 170 175
Tyr Val Gln Phe Arg Glu Phe Asn Leu Lys Asp Pro Pro Leu Asn Pro
180 185 190
<210> 26
<211> 220
<212> PRT
<213> Porcine reproductive and respiratory syndrome virus
<400> 26
Arg His His Phe Thr Pro Ser Glu Arg Gln Leu Cys Leu Ser Ser Ile
1 5 10 15
Gln Thr Ala Phe Asn Gln Gly Ala Gly Thr Cys Ile Leu Ser Asp Ser
20 25 30
Gly Arg Ile Ser Tyr Thr Val Glu Phe Ser Leu Pro Thr His His Thr
35 40 45
Val Arg Leu Ile Arg Val Thr Ala Pro Pro Ser Ala Leu Asp Gln Val
50 55 60
Ile Arg Asn Ala Leu Ala Ser Pro Gly Ser Gly Gly Asp Leu Gly Glu
65 70 75 80
Ala Ile Arg Glu Gln Pro Glu Gln Ala Arg Leu Ala Leu Thr Leu Ala
85 90 95
Ala Ala Glu Ser Glu Arg Phe Val Arg Gln Gly Thr Gly Asn Asp Glu
100 105 110
Ala Gly Ala Ala Asn Ala Asp Val Val Ser Leu Thr Cys Pro Val Ala
115 120 125
Ala Gly Glu Cys Ala Gly Pro Ala Asp Ser Gly Asp Ala Leu Leu Glu
130 135 140
Arg Asn Tyr Pro Thr Gly Ala Glu Phe Leu Gly Asp Gly Gly Asp Val
145 150 155 160
Arg His His Phe Thr Pro Ser Glu Arg Gln Leu Cys Leu Ser Ser Ile
165 170 175
Gln Thr Ala Phe Asn Gln Gly Ala Gly Thr Cys Ile Leu Ser Asp Ser
180 185 190
Gly Arg Ile Ser Tyr Thr Val Glu Phe Ser Leu Pro Thr His His Thr
195 200 205
Val Arg Leu Ile Arg Val Thr Ala Pro Pro Ser Ala
210 215 220
<210> 27
<211> 165
<212> PRT
<213> Porcine reproductive and respiratory syndrome virus
<400> 27
Asn Asn Lys Glu Cys Thr Val Ala Gln Ala Leu Gly Asn Gly Asp Lys
1 5 10 15
Phe Arg Ala Thr Asp Lys Arg Val Val Asp Ser Leu Arg Ala Ile Cys
20 25 30
Ala Asp Leu Glu Gly Ser Ser Ser Pro Leu Pro Lys Val Ala His Asn
35 40 45
Leu Gly Phe Tyr Phe Ser Pro Asp Leu Thr Gln Phe Ala Lys Leu Pro
50 55 60
Ile Glu Leu Asp Pro His Trp Pro Val Val Ser Thr Gln Asn Asn Glu
65 70 75 80
Lys Trp Pro Asp Arg Leu Val Ala Ser Leu Arg Pro Leu Asp Lys Tyr
85 90 95
Ser Arg Ala Cys Ile Gly Ala Gly Tyr Met Val Gly Pro Ser Val Phe
100 105 110
Leu Gly Thr Pro Gly Val Val Ser Tyr Tyr Leu Thr Lys Phe Val Lys
115 120 125
Gly Glu Ala Gln Val Leu Pro Glu Thr Val Phe Ser Thr Gly Arg Ile
130 135 140
Glu Val Asp Cys Arg Glu Tyr Leu Asp Asp Arg Glu Arg Glu Val Ala
145 150 155 160
Ala Ser Leu Pro His
165
<210> 28
<211> 58
<212> PRT
<213> Porcine reproductive and respiratory syndrome virus
<400> 28
Gly Ser Ser Leu Asp Asp Phe Cys Tyr Asp Ser Thr Ala Pro Gln Lys
1 5 10 15
Val Leu Leu Ala Phe Ser Ile Thr Tyr Ala Ser Asn Asp Ser Ser Ser
20 25 30
His Leu Gln Leu Ile Tyr Asn Leu Thr Leu Cys Glu Leu Asn Gly Thr
35 40 45
Asp Trp Leu Ala Asn Lys Phe Asp Trp Ala
50 55
<210> 29
<211> 62
<212> PRT
<213> Porcine reproductive and respiratory syndrome virus
<400> 29
Met Gly Ser Leu Asp Asp Phe Cys Asn Asp Ser Thr Ala Ala Gln Lys
1 5 10 15
Leu Val Leu Ala Phe Ser Ile Thr Tyr Thr Pro Ile Phe Val Ala Gly
20 25 30
Gly Ser Ser Ser Thr Tyr Gln Tyr Ile Tyr Asn Leu Thr Ile Cys Glu
35 40 45
Leu Asn Gly Thr Asp Trp Leu Ser Asn His Phe Asp Trp Ala
50 55 60
<210> 30
<211> 68
<212> PRT
<213> Artificial Sequence
<220>
<223> CD28-PEt
<220>
<221> MISC_FEATURE
<222> (8)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (10)..(11)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (17)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (30)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (32)
<223> Xaa can be any naturally occurring amino acid
<400> 30
Thr Asp Ile Tyr Phe Cys Lys Xaa Glu Xaa Xaa Tyr Pro Pro Pro Tyr
1 5 10 15
Xaa Asp Asn Glu Lys Ser Asn Gly Thr Ile Ile His Arg Xaa Arg Xaa
20 25 30
Lys Arg Gly Trp Glu Gln Leu Glu Gln Cys Gly Tyr Pro Val Gln Arg
35 40 45
Leu Val Ala Leu Tyr Leu Ala Ala Arg Leu Ser Trp Asn Gln Val Asp
50 55 60
Gln Val Ile Arg
65
<210> 31
<211> 28
<212> PRT
<213> Artificial Sequence
<220>
<223> CD28 consensus sequence
<220>
<221> MISC_FEATURE
<222> (8)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (10)..(11)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (17)
<223> Xaa can be any naturally occurring amino acid
<400> 31
Thr Asp Ile Tyr Phe Cys Lys Xaa Glu Xaa Xaa Tyr Pro Pro Pro Tyr
1 5 10 15
Xaa Asp Asn Glu Lys Ser Asn Gly Thr Ile Ile His
20 25
<210> 32
<211> 10
<212> PRT
<213> Artificial Sequence
<220>
<223> CD28 critical region
<220>
<221> MISC_FEATURE
<222> (2)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (4)..(5)
<223> Xaa can be any naturally occurring amino acid
<400> 32
Lys Xaa Glu Xaa Xaa Tyr Pro Pro Pro Tyr
1 5 10
Claims (20)
- 하기를 포함하는 융합 단백질:
(a) 융합 단백질의 N-말단(terminus)에 위치한 항원-제시 세포(antigen-presenting cell; APC)-결합 도메인(domain) 또는 CD91 수용체-결합 도메인;
(b) APC-결합 도메인 또는 CD91 수용체-결합 도메인의 C-말단에 위치한, 하기로 구성된 군으로부터 선별된 단백질 형질도입(transduction) 도메인:
(i) T 세포 민감성 신호-형질도입 펩타이드, 링커, 및 전좌(translocation) 펩타이드를 포함하는 융합 폴리펩타이드(polypeptide), 상기는:
(1) T 세포 민감성 신호-형질도입 펩타이드는 융합 폴리펩타이드의 N-말단에 위치;
(2) T 세포 민감성 신호-형질도입 펩타이드 및 전좌 펩타이드를 연결하는, 서열번호 15를 포함하는 링커; 및
(3) 34 내지 112 아미노산 잔기 길이인 전좌 펩타이드 및 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열을 포함하는 전좌 펩타이드;
(ii) T 세포-민감성 신호-형질도입 펩타이드; 및
(iii) 서열번호 3, 20 또는 4와 최소 90% 동일한 상기 아미노산 서열을 포함하는, 34 내지 112 아미노산 잔기 길이의 전좌 펩타이드; 및
(c) 단백질 형질도입 도메인의 C-말단에 위치한 병원균의 항원, 상기는:
28 내지 53 아미노산 잔기 길이 및 서열번호 31과 최소 90% 동일한 상기 아미노산 서열을 포함하는 T 세포-민감성 신호-형질도입 펩타이드, 상기 Xaa8은 I 또는 L; Xaa10은 V, F 또는 A, Xaa11은 M 또는 L, Xaa17은 L 또는 I; 및
만일 상기 단백질 형질도입 도메인이 전좌 펩타이드(biii)일 때, 수도모나스 엑소톡신A(Pseudomonas exotoxinA; PE) 결합 도메인 I의 아미노산 서열이 없는 APC-결합 도메인 또는 CD91 수용체-결합 도메인.
- 제 1항에 있어서, 상기 APC-결합 도메인 또는 CD91 수용체-결합 도메인은 서열번호 5, 9, 6, 7, 및 8로 구성되는 군으로부터 선택된 서열과 최소 90% 동일한 아미노산 서열을 포함하는 폴리펩타이드인 융합 단백질.
- 제 1항에 있어서, 추가적으로 융합 단백질의 C-말단에 위치한 골지체(endoplasmic reticulum) 유지 서열을 포함하는 융합 단백질.
- 제 3항에 있어서, 상기 골지체 유지 서열은 Lys-Asp-Glu-Leu(서열번호 14)의 아미노산 서열을 포함하는 융합 단백질.
- 제 1항에 있어서, 상기 융합 단백질은 만약 항원이 10 또는 그 이상의 항원결정기(epitope)를 갖는다면 C-말단에 골지체 유지 서열이 없는 융합 단백질.
- 제 1항에 있어서, 상기 단백질 형질도입 도메인은 융합 폴리펩타이드인 융합 단백질(bi).
- 제 1항에 있어서, 상기 단백질 형질도입 도메인은 T 세포-민감성 신호-형질도입 펩타이드인 융합 단백질(bii).
- 제 7항에 있어서, 추가적으로 상기 단백질 형질도입 도메인 및 항원 사이의, 서열번호 15를 포함하는 추가적인 링커를 포함하는 융합 단백질.
- 제 1항에 있어서, 상기 단백질 형질도입 도메인은 전좌 펩타이드인 융합 단백질(biii).
- 제 9항에 있어서, 추가적으로 APC-결합 도메인 또는 CD91 수용체-결합 도메인 사이의 서열번호 15를 포함하는 추가적인 링커 및 상기 형질도입 펩타이드를 포함하는 융합 단백질.
- 제 1항에 있어서, 상기 형질도입 도메인은 서열번호 30의 서열을 포함하는 융합 단백질.
- 하기를 포함하는 백신 조성물(vaccine composition):
(a) 치료적으로 효과적인 양의 제 1항의 융합 단백질; 및
(b) 아주번트(adjuvant).
- 제 1항에 있어서, 상기 APC-결합 도메인 또는 CD91 수용체-결합 도메인은 서열번호 5, 9, 6, 7 및 8로 구성되는 군으로부터 선택된 아미노산 서열을 포함하는 폴리펩타이드인 융합 단백질.
- 제 1항에 있어서, 상기 T 세포 민감성 신호-형질도입 펩타이드는 서열번호 1 또는 2와 최소 90% 동일한 아미노산 서열을 포함하는 융합 단백질.
- 제 14항에 있어서, 상기 T 세포 민감성 신호-형질도입 펩타이드는 서열번호 1 및 2로 구성되는 군으로부터 선택된 아미노산 서열을 포함하는 융합 단백질.
- 제 1항에 있어서, 상기 형질도입 펩타이드는 서열번호 3, 20, 및 4로 구성되는 군으로부터 선택되는 아미노산 서열을 포함하는 융합 단백질.
- 강화된 병원균 항원-특이적인 T 세포 반응의 유도에 사용하기 위한 상기 항 중 어느 하나에서 청구된 것으로서 융합 단백질 또는 백신 조성물.
- 질병 세포의 세포 막(membrane)에 있는 클래스 I MHC 분자(class I MHC molecule)를 통해 항원을 나타내는 질병 세포를 사멸에 사용하기 위한 상기 항 중 어느 하나에서 청구된 것으로서 융합 단백질 또는 백신 조성물.
- 제 18항에서 있어서, 질병 세포가 암 세포 일 때 사용하기 위한 융합 단백질 또는 백신 조성물.
- 병원균에 의해 야기되는 감염의 예방, 치료 및/또는 감염에 의해서 야기되는 증상의 최소화에 사용하기 위한 상기 청구항 중 어느 하나에서 청구된 것으로 융합 단백질 또는 백신 조성물.
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