KR20230004706A - 폐부종의 치료 또는 예방용 rock 억제제 - Google Patents
폐부종의 치료 또는 예방용 rock 억제제 Download PDFInfo
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- Investigating Or Analysing Biological Materials (AREA)
Abstract
본 발명은 바이러스 감염과 관련된 폐부종의 치료 또는 예방에 사용하기 위한 ROCK 억제제에 관한 것이다. 본 발명은 추가로 폐 상피 세포에서 정단 나트륨-칼륨-ATPase(NKA) 국소화를 방지하거나 감소시키는 데 있어서 억제제의 효과의 결정 또는 시험관내 시험 시스템의 용도에 관한 것이다. 또한, 폐부종의 예방 및/또는 치료에 효과적인 분자를 검출하기 위한 방법이 제공된다. 마지막으로, 본 발명은 시험 시스템에 관한 것이다.
Description
발명의 기술 분야
본 발명은 바이러스 감염과 관련된 폐부종의 치료 또는 예방에 사용하기 위한 ROCK 억제제에 관한 것이다. 본 발명은 추가로 폐 상피 세포에서 정단 나트륨-칼륨-ATPase(sodium-potassium-ATPase; NKA) 국소화를 방지하거나 감소시키는 데 있어서 억제제의 효과의 결정 또는 시험관내 시험 시스템의 용도에 관한 것이다. 또한, 폐부종의 예방 및/또는 치료에 효과적인 분자를 검출하기 위한 방법이 제공된다. 마지막으로, 본 발명은 시험 시스템에 관한 것이다.
설명
폐부종은 다수의 상이한 요인에 의해 초래될 수 있다. 이는 심장성 폐부종으로 지칭되는 심부전과 관련되거나, 비-심장성 폐부종으로 지칭되는 바이러스 감염과 같은 다른 원인과 관련될 수 있다. 예를 들어, 인간의 인플루엔자 A 바이러스(Influenza A Virus; IAV) 감염은 폐 손상, 및 폐포 폐강에 체액의 과도한 축적(폐부종)에 의해 초래되는 "급성 호흡 곤란 증후군(Acute Respiratory Distress Syndrome)"(ARDS)으로 이어질 수 있고, 치료하지 않으면 저산소혈증 및 사망으로 이어질 수 있다.
효과적인 치료를 위해서는 신속한 진단과 조기 개입이 필요하다. 결과적으로, 지난 2세기 동안 폐부종을 신속하게 진단하고 치료에 대한 반응을 추적하기 위한 임상 도구를 개발하기 위한 집중적인 노력이 이루어졌다. 이러한 도구의 이상적인 특성은 높은 민감도 및 특이성, 용이한 이용 가능성, 및 완전한 임상 발현의 발생 전에 폐수(lung water)의 조기 축적을 진단하는 능력을 포함할 것이다. 또한, 임상의는 혈관외 폐수 축적을 정확하게 정량화하고 폐부종의 고 투과성 병인으로부터 정수압을 구별하는 능력을 높이 평가한다.
따라서, 폐부종의 요법에 대한 당 분야의 요구가 여전히 존재한다.
본 발명의 해결책이 하기에 기술되고, 실시예에서 예증되며, 도면에 예시되고 청구범위에 반영된다.
본 발명은
i)
폐 상피 세포에서 정단 NKA 국소화를 방지하거나,
ii)
ROCK 억제제의 투여 전 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킴으로써,
폐부종의 치료 또는 예방에 사용하기 위한 ROCK 억제제로서,
폐부종은 바이러스 감염과 관련되고, 바이러스는 아르티큘라바이러스목(Articulavirales)(예를 들어, 오르토믹소바이러스과(Orthomyxoviridae)), 모노네가바이러스목(Mononegavirales)(예를 들어, 뉴모바이러스과(Pneumoviridae)) 및/또는 분야바이러스목(Bunyavirales)(예를 들어, 한타바이러스과(Hantaviridae))인, ROCK 억제제에 관한 것이다.
본 발명은 또한
i)
폐 상피 세포에서 정단 NKA 국소화를 방지하거나,
ii)
ROCK 억제제의 투여 전 존재하는 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킴으로써,
폐부종의 예방 및/또는 치료 방법에 사용하기 위한 ROCK 억제제를 포함하는 조성물로서,
폐부종은 바이러스 감염과 관련되고, 바이러스는 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목인, 조성물에 관한 것이다.
본 발명은 또한 폐 상피 세포에서의 바이러스-유도 정단 NKA 국소화를 방지하거나 감소시키는 데 효과적인 억제제의 결정을 위한 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템의 용도에 관한 것이다.
또한, 본 발명은 폐부종의 예방 및/또는 치료에 효과적인 분자를 검출하는 방법으로서, 관심 화합물과 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템을 접촉시키는 것을 포함하고, 관심 화합물은 접촉 전의 시험관내 시험 시스템과 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시키는, 방법에 관한 것이다.
본 발명은 또한 폐 상피 세포에서 RNA 바이러스 관련 정단 NKA 국소화를 예방함으로써 폐부종을 갖거나 이의 위험이 있는 대상체를 치료하는 방법에 관한 것이다.
본 발명은 또한 i) ROCK 억제제; ii) 폐 상피 세포 iii) 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스; 및 iv) NKA의 검출을 위한 수단을 포함하는 시험 시스템에 관한 것이다.
도면은 하기를 나타낸다:
도 1: ROCK 억제는 정단 나트륨-칼륨-ATPase(NKA) 국소화를 방지한다. 대조: 비감염 Calu3 세포에서 NKA(연회색)의 기저측 분포. PR8: 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)(20 h p.i.)에 감염된 Calu3 세포에서의 정단 NKA 분포. PR8+Rho XIII: 인플루엔자 바이러스 A/푸에르토리코/8/34에 감염되고 ROCK 억제제로 처리된(감염 후(p.i.) 20 h) Calu3 세포에서의 NKA 분포. (NKAα-(진회색), 바이러스 핵단백질(NP)(연회색), 핵(진회색)). 섹션 두께: 0.25 μm, 스케일 바: 10 μm.
도 2: ROCK 억제는 분극화된(감염) Calu3 세포에서 벡터 물 수송을 개선한다. 모의: 미처리 및 비감염 분극화된 Calu3 세포에 대한 벡터 물 수송(VWT)을 측정하고 100%로 설정하였다. PR8: 비처리 및 인플루엔자 바이러스 A/푸에르토리코/8/34 (H1N1, MOI = 2)-감염 Calu3 세포를 통한 VWT. Rho XIII + PR8: 처리(ROCK 억제제: XIII, 5 μM) 및 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)-감염 Calu3 세포를 통한 VWT. 아밀로라이드: 아밀로라이드(ENaC 차단)-처리 및 비감염 Calu3 세포(음성 대조, ENaC-의존성 부분을 나타냄)를 통한 VWT. A) 8 시간 p.i., B) 16 시간 p.i.
도 3: ROCK 억제는 비감염 및 감염 Calu3 세포의 세포 생존율을 증가시킨다. 모의: Calu3 세포를 0, 6, 16, 24 시간 동안 인큐베이션하고 생존율을 결정하고(MTT 시험) 100%로 설정하였다. Rho XIII: Calu3 세포를 ROCK 억제제 Rho XIII(5 μM)로 0, 6, 16, 24 시간 동안 처리한 후 생존율을 결정하였다. H1N1: Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)(MOI = 2)에 감염시키고 생존율을 결정하였다. Rho XIII+H1N1: Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)에 감염시키고, ROCK 억제제 Rho XIII(5 μM)로 0, 6, 16, 24 시간 동안 처리한 후, 생존율을 결정하였다.
도 4: ROCK 억제는 시험관내에서 바이러스-유도 상피 세포 손상을 감소시킨다. 모의: 전면으로 생장되고 분극화된 모의-감염 Calu3 세포를 쿠마시(Coomassie)(청색)로 염색하였다. H1N1: 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(MOI = 5)에 감염시키고 24 h p.i.에 쿠마시로 염색하였다. Rho XIII: 전면으로 생장된 Calu3 세포를 ROCK 억제제(Rho XIII, 5 μM)로 처리하고, 24 시간 후에 쿠마시로 염색하였다. H1N1+Rho XIII: 전면으로 생장된 Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34에 감염시키고, ROCK 억제제(Rho XIII, 5 μM)로 처리하고, 24 h p.i.에 쿠마시로 염색하였다.
도 5: ROCK 억제는 감염된 C57BL/6 마우스의 체중 감소의 경향적 감소를 야기한다. 파수딜(Fasudil) HCL: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 1일 1회 처리된 비감염 마우스로부터의 체중. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스의 체중. PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)으로 1일 1회 처리된 마우스의 체중. 실험은 7일 동안만 계획되었다. 이후, 윤리적인 이유로 마우스를 안락사시켰다. 따라서, 흑색 및 적색 곡선은 서로 훨씬 더 강하게(유의하게) 분리될 수 있다.
도 6: ROCK 억제는 감염된 마우스 폐의 체액 중량(습윤 건조 중량 비율)을 감소시킨다. 모의: 비처리된 모의-감염 마우스의 습윤-건조 폐 비율. 파수딜 HCL: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 6일 동안 1일 1회 처리된 마우스의 폐의 습윤-건조 비율. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스의 습윤-건조 폐 비율(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 마우스의 습윤-건조 폐 비율(7일차 p.i.).
도 7: ROCK 억제는 폐의 조직 구조를 개선하고 감염된 마우스의 세포 폐포 침윤을 감소시킨다. 대조: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 6일 동안 1일 1회 처리된 동물을 나타내는 폐 조직 절편(다양한 확대)(7일차 p.i.). PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 동물을 나타내는 폐 조직 절편(다양한 확대)(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 동물을 나타내는 폐 조직 절편(다양한 배율)(7일차 p.i.)으로서, 하나의 개략적 이미지(4x) 및 선택된 절편의 2배 확대(10x, 20x)를 보여줌.
도 8: ROCK 억제는 침윤물의 수를 감소시킨다(정량화). 대조: 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 5 마리의 마우스로부터의 폐 조직 절편(다양한 배율)(7일차 p.i.). PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 5 마리의 마우스로부터의 폐 조직 절편(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 5 마리의 마우스로부터의 폐 조직 절편(7일차 p.i.). 폐 조직 절편을 Aperio CS2 스캐너(Leica Biosystems Imaging Inc., CA, USA)로 디지털화시키고, "Aperio v9 핵 계수 알고리즘" 소프트웨어(Leica Biosystems Imaging Inc., CA, USA) 및 결정된 각 그룹의 폐 절편의 평균 면적(평균 표면/mm2), 평균 총 세포 수/표면 및 계산된 mm2 당 평균 세포 수(총 세포 수/mm2)을 사용하여 분석하였다.
도 9: ROCK 억제는 감염된 마우스의 폐에서 바이러스 역가를 감소시킨다. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스로부터의 폐 균질화물의 바이러스 역가(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 마우스로부터의 폐 균질화물의 바이러스 역가(7일차 p.i.). n = 3
도 10: PR8-감염 세포에서 NKA, M2 및 HA의 세포 표면 발현. PR8-감염 Calu3 세포 (+/-) ROCK 억제제 처리의 세포 표면 상의 NKA, M2 및 HA의 양은 NKA에 특이적인 항체(베타-서브유닛, HA 및 M2(n=16))를 사용하여 비-투과성 세포의 온-셀 웨스턴 블롯(On-Cell Western Blot) 분석에 의해 평가하였다.
도 11: 상이한 IAV 서브타입에 감염된 Calu3 세포의 원형질막 내의 NKAβ1의 재분포.
(A) Calu3 세포에서 상이한 IAV 서브타입의 생장. 세포는 MOI 0.01에서 지시된 바이러스에 감염되었고, 바이러스 역가는 지시된 시점에 병소 검정에 의해 결정되었다. 막대 그래프는 평균 + SD, n=3을 나타낸다.
(B) 상이한 IAV 균주에 감염된 Calu3 세포의 정단막에 대한 NKAβ1의 정량화. 96-웰 플레이트 상에서 Calu3 세포의 단층은 지시된 바이러스(MOI:2)에 감염되었고 20 h p.i.이다. OCWB 분석이 수행되었다. 데이터는 평균+SD, n=16을 나타낸다.
도 1: ROCK 억제는 정단 나트륨-칼륨-ATPase(NKA) 국소화를 방지한다. 대조: 비감염 Calu3 세포에서 NKA(연회색)의 기저측 분포. PR8: 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)(20 h p.i.)에 감염된 Calu3 세포에서의 정단 NKA 분포. PR8+Rho XIII: 인플루엔자 바이러스 A/푸에르토리코/8/34에 감염되고 ROCK 억제제로 처리된(감염 후(p.i.) 20 h) Calu3 세포에서의 NKA 분포. (NKAα-(진회색), 바이러스 핵단백질(NP)(연회색), 핵(진회색)). 섹션 두께: 0.25 μm, 스케일 바: 10 μm.
도 2: ROCK 억제는 분극화된(감염) Calu3 세포에서 벡터 물 수송을 개선한다. 모의: 미처리 및 비감염 분극화된 Calu3 세포에 대한 벡터 물 수송(VWT)을 측정하고 100%로 설정하였다. PR8: 비처리 및 인플루엔자 바이러스 A/푸에르토리코/8/34 (H1N1, MOI = 2)-감염 Calu3 세포를 통한 VWT. Rho XIII + PR8: 처리(ROCK 억제제: XIII, 5 μM) 및 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)-감염 Calu3 세포를 통한 VWT. 아밀로라이드: 아밀로라이드(ENaC 차단)-처리 및 비감염 Calu3 세포(음성 대조, ENaC-의존성 부분을 나타냄)를 통한 VWT. A) 8 시간 p.i., B) 16 시간 p.i.
도 3: ROCK 억제는 비감염 및 감염 Calu3 세포의 세포 생존율을 증가시킨다. 모의: Calu3 세포를 0, 6, 16, 24 시간 동안 인큐베이션하고 생존율을 결정하고(MTT 시험) 100%로 설정하였다. Rho XIII: Calu3 세포를 ROCK 억제제 Rho XIII(5 μM)로 0, 6, 16, 24 시간 동안 처리한 후 생존율을 결정하였다. H1N1: Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)(MOI = 2)에 감염시키고 생존율을 결정하였다. Rho XIII+H1N1: Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)에 감염시키고, ROCK 억제제 Rho XIII(5 μM)로 0, 6, 16, 24 시간 동안 처리한 후, 생존율을 결정하였다.
도 4: ROCK 억제는 시험관내에서 바이러스-유도 상피 세포 손상을 감소시킨다. 모의: 전면으로 생장되고 분극화된 모의-감염 Calu3 세포를 쿠마시(Coomassie)(청색)로 염색하였다. H1N1: 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34(MOI = 5)에 감염시키고 24 h p.i.에 쿠마시로 염색하였다. Rho XIII: 전면으로 생장된 Calu3 세포를 ROCK 억제제(Rho XIII, 5 μM)로 처리하고, 24 시간 후에 쿠마시로 염색하였다. H1N1+Rho XIII: 전면으로 생장된 Calu3 세포를 인플루엔자 바이러스 A/푸에르토리코/8/34에 감염시키고, ROCK 억제제(Rho XIII, 5 μM)로 처리하고, 24 h p.i.에 쿠마시로 염색하였다.
도 5: ROCK 억제는 감염된 C57BL/6 마우스의 체중 감소의 경향적 감소를 야기한다. 파수딜(Fasudil) HCL: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 1일 1회 처리된 비감염 마우스로부터의 체중. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스의 체중. PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)으로 1일 1회 처리된 마우스의 체중. 실험은 7일 동안만 계획되었다. 이후, 윤리적인 이유로 마우스를 안락사시켰다. 따라서, 흑색 및 적색 곡선은 서로 훨씬 더 강하게(유의하게) 분리될 수 있다.
도 6: ROCK 억제는 감염된 마우스 폐의 체액 중량(습윤 건조 중량 비율)을 감소시킨다. 모의: 비처리된 모의-감염 마우스의 습윤-건조 폐 비율. 파수딜 HCL: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 6일 동안 1일 1회 처리된 마우스의 폐의 습윤-건조 비율. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스의 습윤-건조 폐 비율(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 마우스의 습윤-건조 폐 비율(7일차 p.i.).
도 7: ROCK 억제는 폐의 조직 구조를 개선하고 감염된 마우스의 세포 폐포 침윤을 감소시킨다. 대조: 파수딜 HCl(염수 용액 중 10 mg/kg)으로 6일 동안 1일 1회 처리된 동물을 나타내는 폐 조직 절편(다양한 확대)(7일차 p.i.). PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 동물을 나타내는 폐 조직 절편(다양한 확대)(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 동물을 나타내는 폐 조직 절편(다양한 배율)(7일차 p.i.)으로서, 하나의 개략적 이미지(4x) 및 선택된 절편의 2배 확대(10x, 20x)를 보여줌.
도 8: ROCK 억제는 침윤물의 수를 감소시킨다(정량화). 대조: 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 5 마리의 마우스로부터의 폐 조직 절편(다양한 배율)(7일차 p.i.). PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 5 마리의 마우스로부터의 폐 조직 절편(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 5 마리의 마우스로부터의 폐 조직 절편(7일차 p.i.). 폐 조직 절편을 Aperio CS2 스캐너(Leica Biosystems Imaging Inc., CA, USA)로 디지털화시키고, "Aperio v9 핵 계수 알고리즘" 소프트웨어(Leica Biosystems Imaging Inc., CA, USA) 및 결정된 각 그룹의 폐 절편의 평균 면적(평균 표면/mm2), 평균 총 세포 수/표면 및 계산된 mm2 당 평균 세포 수(총 세포 수/mm2)을 사용하여 분석하였다.
도 9: ROCK 억제는 감염된 마우스의 폐에서 바이러스 역가를 감소시킨다. PR8: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염된 마우스로부터의 폐 균질화물의 바이러스 역가(7일차 p.i.). PR8 + 파수딜 HCL: IAV A/푸에르토리코/8/32(H1N1, 500 PFU/마우스)에 감염되고 파수딜 HCl(염수 용액 중 10 mg/kg)로 6일 동안 1일 1회 처리된 마우스로부터의 폐 균질화물의 바이러스 역가(7일차 p.i.). n = 3
도 10: PR8-감염 세포에서 NKA, M2 및 HA의 세포 표면 발현. PR8-감염 Calu3 세포 (+/-) ROCK 억제제 처리의 세포 표면 상의 NKA, M2 및 HA의 양은 NKA에 특이적인 항체(베타-서브유닛, HA 및 M2(n=16))를 사용하여 비-투과성 세포의 온-셀 웨스턴 블롯(On-Cell Western Blot) 분석에 의해 평가하였다.
도 11: 상이한 IAV 서브타입에 감염된 Calu3 세포의 원형질막 내의 NKAβ1의 재분포.
(A) Calu3 세포에서 상이한 IAV 서브타입의 생장. 세포는 MOI 0.01에서 지시된 바이러스에 감염되었고, 바이러스 역가는 지시된 시점에 병소 검정에 의해 결정되었다. 막대 그래프는 평균 + SD, n=3을 나타낸다.
(B) 상이한 IAV 균주에 감염된 Calu3 세포의 정단막에 대한 NKAβ1의 정량화. 96-웰 플레이트 상에서 Calu3 세포의 단층은 지시된 바이러스(MOI:2)에 감염되었고 20 h p.i.이다. OCWB 분석이 수행되었다. 데이터는 평균+SD, n=16을 나타낸다.
건강한/대조 폐 상피 세포에서, NKA는 본질적으로 기저측으로 국소화된다. 이러한 상황에서, 나트륨은, 예를 들어, 정단 국소화된 나트륨 채널(ENaC)을 통해 폐 상피 세포로 흡수된다. NKA는 이후 폐 상피 세포로부터 상피하 간질 공간으로 나트륨을 내보낸다. 물은 아쿠아포린을 통해 및 다른 세포내 경로를 통해 상피하 간질 공간으로 이러한 나트륨 농도구배를 따르게 된다. 이러한 방식으로, 폐부종의 발병이 예방된다. 따라서, 결과적으로 NKA는 기능적으로 손상된 경우 부종 제거의 주요 제한 인자를 나타낸다(Peteranderl et al., (2019) "Influenza A virus infection induces apical redistribution of Na+, K+-ATPase in lung epithelial cells in vitro and in vivo" American Journal of Respiratory Cell and Molecular Biology Volume 61 Number 3, pp. 395-397).
구체적으로, 본 발명자들이 발견한 것은, 아르티큘라바이러스목 및 이에 따라 또한 아마도 모노네가바이러스목 및/또는 분야바이러스목의 인플루엔자 A 바이러스(IAV)로의 폐 상피 세포, 바람직하게는 폐포 상피 세포(예를 들어, Calu3 세포)의 바이러스 감염 시 NKA는 폐 상피 세포의 기저측 쪽에서 정단 쪽으로 적어도 부분적으로 재분포된다는 것이다. 이러한 재분포로 인해, 나트륨 농도구배뿐만 아니라 정단에서 기저측 폐 세포 쪽으로의 방향성 물 수송이 방해를 받는다. 이는 폐부종의 발병으로 이어진다. 특히, 밀착 접합 복합체는 이러한 바이러스 감염에 의해 영향을 받지 않는데, 이는 세포 극성이 유지된다는 것을 지시한다(Peteranderl et al., (2019) "Influenza A virus infection induces apical redistribution of Na+, K+-ATPase in lung epithelial cells in vitro and in vivo" American Journal of Respiratory Cell and Molecular Biology Volume 61 Number 3, pp. 395-397).
놀랍게도, ROCK 억제제는 (i) 바이러스-감염된 인간 극성 폐 상피 세포(Calu3 세포와 같이)의 기저측 쪽에서 정단 쪽으로의 NKA의 IAV-유도 병리학적 재분포를 방지하고, (ii) 세포 배양에서 지시된 체액 수송(정단에서 기저측으로)을 회복시키는 것으로 밝혀졌다. 따라서, ROCK 억제제는 IAV-감염 폐 상피 세포에서 세포내 물 수송에 직접 영향을 미침으로써 폐부종을 치료 및/또는 예방한다. 특히, 치료 또는 예방은 IAV-유도 정단 NKA 국소화의 방지를 통해 나트륨 농도구배를 회복시킴으로써 정단에서 기저측 쪽으로의 세포내 지시된 물 수송을 장내 공간으로 재설정함으로써 달성된다. 이론에 국한되지 않으면서, 본 발명자들은 ROCK 억제제가 NKA의 정단 쪽으로의 재분포를 방지할 뿐만 아니라 기저측으로 NKA를 안정화시키는 것으로 사료한다.
또한, ROCK 억제제는 (iii) 세포 배양에서 IAV 증식의 세포변성 효과(CPE)를 크게 감소시키는 것으로 밝혀졌다. 또한, 동물 실험은 ROCK 억제가 (iv) 감염된 마우스의 체중 저하의 감소를 야기하고, (v) 감염된 마우스 폐의 체액 중량을 감소시키고, (vi) 폐의 조직 구조를 안정화시키고, (vii) 세포 폐포 침윤을 감소시키고, (viii) 감염된 마우스의 폐에서 바이러스 역가를 감소시킨다는 것을 나타냈다.
따라서, 본 발명은
i)
폐 상피 세포에서 정단 NKA 국소화를 방지하거나,
ii)
ROCK 억제제의 투여 전 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킴으로써,
폐부종의 치료 또는 예방에 사용하기 위한 ROCK 억제제로서,
폐부종은 바이러스 감염과 관련되고, 바이러스는 아르티큘라바이러스목(예를 들어, 오르토믹소바이러스과), 모노네가바이러스목(예를 들어, 뉴모바이러스과) 및/또는 분야바이러스목(예를 들어, 한타바이러스과)인, ROCK 억제제에 관한 것이다.
본원에서 사용되는 "ROCK 억제제"는 임의의 적합한 ROCK 억제제일 수 있다. 일반적으로, ROCK 억제제는 Rho-관련 단백질 키나제(ROCK) 경로의 억제제이다. ROCK 경로는 당업자에게 공지되어 있으며, 특히 문헌[Liao et al. (2007) "Rho Kinase (ROCK) Inhibitors" J. Cardiovasc Pharmacol. 50(1):17-24 and Amano et al. (2010) "Rho-Kinase/ROCK: A Key Regulator of the Cytoskeleton and Cell Polarity" Cytoskeleton (Hoboken). 2010 Sep; 67(9): 545-554]에 기재되어 있다. 따라서, 용어 "ROCK 신호전달 경로"는 (활성) Rho-관련 키나제(Rho-키나제/ROCK/ROK)에 의해 개시되는 세포 사건의 캐스케이드를 지칭한다.
세포 사건의 캐스케이드는, 예를 들어, Rho로부터 시작할 수 있다. Rho 서브패밀리는 Ras 패밀리에서 소분자 G 단백질의 구성원이고 GTPase 활성을 갖는다. 따라서, Rho는 활성화된 상태(Rho-GTP)와 비활성 상태(Rho-GDP) 사이에서 전환된다. 예를 들어, 리소포스파트산(LPA) 또는 스핑고신-1 포스페이트(S1P)를 통한 Rho의 자극 시, GTP-결합된 Rho(활성 Rho)가 생성된다. Rho의 하류에서 하나의 이펙터 분자는 Rho-관련 키나제(Rho-키나제/ROCK/ROK)이다. 따라서, Rho-GTP는 ROCK를 활성화시킬 수 있다. 그러나, ROCK는 또한 Rho와 독립적으로, 즉, 예를 들어, 아미노-말단 트랜스인산화를 통해 활성화될 수 있다. 활성화 시 ROCK 단백질은 F-액틴과 같은 많은 하류 표적을 인산화시킨다.
다른 한 편으로, ROCK는 Gem 및 Rad와 같은 다른 작은 GTP-결합 단백질에 의해 억제될 수 있다.
구체적으로, ROCK는 또한 문헌[Liao et al. (2007) "Rho Kinase (ROCK) Inhibitors" J. cardiovasc Pharmacol. 50(1):17-24]에 기재된 바와 같이 아미노-말단 단백질 세린/트레오닌 키나제 도메인, 및 이어서 Rho-결합 도메인(RBD)을 함유하는 중간 코일-코일-형성 영역, 및 플렉스트린 상동성(PH) 모티프 내에 위치한 카르복시-말단 시스테인-부유 도메인(CRD)으로 이루어진다. 지금까지 2개의 ROCK 이소형, 즉, ROCK1 및 ROCK2가 확인되었다.
본원에서 사용되는 ROCK의 "억제제"는 ROCK의 활성 또는 ROCK 경로를 감소시키거나 억제할 수 있는 임의의 적합한 억제제로서 정의된다. 억제제는 ROCK의 활성 또는 ROCK 경로를 감소시키거나 없애는 화합물/분자일 수 있다. 억제제는 ROCK를 인코딩하는 유전자의 전사를 감소 또는 차단하고/차단하거나 ROCK를 인코딩하는 mRNA의 번역을 감소시킴으로써 이러한 효과를 달성할 수 있다. 또한, 억제제는 ROCK(들)이 억제제의 부재 하에서보다 억제제의 존재 하에서 감소된 효율로 이의 생화학적 기능을 수행하도록 할 수 있다. 또한, 억제제는 ROCK가 억제제의 부재 하에서보다 활성화제의 존재 하에서 감소된 효율로 이의 세포 기능을 수행하게 할 수 있다.
따라서, 용어 "억제제"는 또한 ROCK 경로, 예를 들어, Gem 및 Rad에 직접적으로 감소시키는 효과를 갖는 분자/화합물뿐만 아니라, 예를 들어, ROCK 경로를 양으로 조절하는(예를 들어, Rho-GTP, LPA 또는 S1P와 같이 활성화시키는) 분자와 상호작용함으로써 간접적으로 감소시키는 분자를 포함한다.
억제제는 또한 억제될 경로의 길항제일 수 있다.
화합물/분자가 ROCK의 활성 또는 ROCK 경로를 감소시키거나 억제할 수 있는지 시험하는 방법은 당업자에게 공지되어 있다. 예를 들어, ROCK 경로 또는 ROCK의 억제제는 당업자에게 공지된 표준 시험을 수행함으로써 시험될 수 있다. 당업자는 탐침자를 미오신 포스페이트 표적 서브유닛 1(MYPT1)과 접촉시킬 수 있다. MYPT1은 트레오닌 696(T696)에서 이러한 기질을 인산화하는 ROCK1 및 ROCK2의 기질이다. 이러한 인산화는 항-포스포-MYPT1-Thr696 항체에 의해 검출될 수 있다. 재조합 활성 ROCK2는 양성 대조로서 사용될 수 있다. 이러한 시험은 상업적으로 이용 가능하며, 예를 들어, abcam(ROCK 활성 검정; abcam Cat# ab211175) 또는 Millipore(Millipore Cat# CSA001)로부터 입수될 수 있다.
억제제는 ROCK 경로 또는 ROCK 활성을 억제제의 첨가 없이 또는 첨가 전의 ROCK 경로 또는 ROCK 활성과 비교하여 10%, 20%, 30%, 40%, 50%, 60%, 70%, 80%, 90%, 100% 이상까지 ROCK 경로 또는 ROCK 활성을 억제하거나 감소시킬 수 있다.
ROCK 억제제는 소분자, 화합물, 항체와 같은 결합 분자, siRNA와 같은 핵산 분자 또는 전구약물일 수 있다.
본원에서 사용되는 "소분자"는 ROCK 경로(Rho/ROCK 경로) 또는 ROCK 활성을 감소/억제할 수 있는 임의의 소분자일 수 있다. 소분자는 저분자량의 유기 화합물일 수 있다. 저분자량은 화합물이 900 달톤(da) 미만, 800 da 미만, 700 da 미만, 600 da 미만 또는 500 da 미만의 중량을 갖는 것을 의미할 수 있다. 예를 들어, 소분자는 약 300 da의 분자량을 가질 수 있다. 소분자의 크기는 당 분야에 널리 공지된 방법, 예를 들어, 질량 분광법에 의해 결정될 수 있다. 예시적인 소분자 ROCK 억제제는 Y-27632 및 CCG-1423뿐만 아니라 파수딜 HCl(327.830898 Da), RKI-1447(326.374812 Da) 및 하이드록시파수딜(307.369824 Da)을 포함한다.
본원에서 사용되는 "화합물"은 ROCK 경로(Rho/ROCK 경로) 또는 ROCK를 감소/억제할 수 있는 임의의 화합물일 수 있다. 억제제로서 사용될 수 있는 화합물/분자는 각각의 경로/ROCK를 억제 또는 감소시킬 수 있거나 억제될 경로/ROCK의 저해제를 활성화시키거나 억제될 경로/ROCK의 활성화제를 억제하는 임의의 화합물/분자일 수 있다.
본원에서 사용되는 "결합 분자"는 ROCK 경로(Rho/ROCK 경로) 또는 ROCK를 감소/억제할 수 있는 임의의 결합 분자일 수 있다. 예를 들어, 결합 분자는 항체, 항체 단편 또는 상이한 특이성을 갖는 2개의 결합 부위를 포함하는 2가 항체 단편일 수 있다.
항체는 임의의 항-ROCK1 및/또는 항-ROCK2 항체일 수 있다. 이러한 항체는 상업적으로 이용 가능하다. 예를 들어, 항체는 abcam의 항-ROCK1 항체 [EP786Y] 또는 [EPR638Y](ab45171 또는 ab230799), abcam의 항-ROCK2 항체(ab71598), Santa Cruz의 Rock1 항체(B-1)(sc-374388), Cell Signaling로부터의 항-ROCK1 항체(C8F7) mAb #4035, Cell Signaling으로부터의 항-ROCK2 항체(D1B1) mAb #9029 또는 Cell Signaling로부터의 항-ROCK2 항체 #8236일 수 있다.
이러한 2가 항체 단편의 비제한적인 예는 (Fab)2'-단편, 2가 단쇄 Fv 단편, bsFc-1/2-이량체 또는 bsFc-CH3-1/2 이량체를 포함한다. 결합 분자는 또한 단일 결합 부위만을 가질 수 있고, 즉, 1가일 수 있다. 1가 결합 분자의 예는 1가 항체 단편, 항체-유사 결합 특성을 갖는 단백질성 결합 분자를 포함하나, 이에 제한되지 않는다. 1가 항체 단편의 예는 Fab 단편, Fv 단편, 단쇄 Fv 단편(scFv) 또는 scFv-Fc 단편을 포함하나, 이에 제한되지 않는다.
대안적으로, 결합 분자는 또한 "듀오칼린"으로도 알려진 리포칼린 뮤테인과 같은 2가 단백질성 인공 결합 분자일 수 있다.
결합 분자는 또한 항체-유사 결합 특성을 갖는 단백질성 결합 분자일 수 있다. 예시적이지만 비제한적인 단백질성 결합 분자는 압타머, 리포칼린 패밀리의 폴리펩타이드에 기반한 뮤테인, 글루바디, 안키린 스캐폴드에 기반한 단백질, 결정질 스캐폴드에 기반한 단백질, 애드넥틴, 아비머 또는 (재조합) 수용체 단백질을 포함한다.
억제제는 또한 핵산 분자일 수 있다. 예는 ROCK 경로(Rho/ROCK 경로) 또는 ROCK를 억제할 수 있는 RNA, siRNA, miRNA 또는 비-단백질성 압타머를 포함한다. 이러한 압타머는 특정 표적 분자에 결합하는 올리고핵산이다. 이러한 압타머는 DNA 또는 RNA 압타머로 분류될 수 있다. 이들은 (대개 짧은) 올리고뉴클레오타이드 가닥으로 이루어진다. 또한, 핵산 분자는 억제될 경로의 활성화제, 프로모터 또는 인핸서를 저해하는 데 사용될 수 있다.
ROCK1 및 ROCK2에 대한 siRNA는 당업자에게 상업적으로 이용 가능하다. 예를 들어, siRNA는 Santa Cruz의 Rock-1 siRNA(h)(sc-76025), Santa Cruz의 ROCK2 siRNA(h)(sc-29474), Thermos Fischer Scientific의 ROCK-1 siRNA(h) #4390824 또는 Thermo Fischer Scientific의 ROCK-2 siRNA(h) AM51331일 수 있다.
억제제는 또한 ROCK 경로(Rho/ROCK 경로)/ROCK를 억제할 수 있는 전구약물일 수 있다. "전구약물"은 약리학적으로 본질적으로 불활성이고, 전구약물과 함께 투여된 대상체의 체내에서 이의 활성 형태로 대사된다. 적합한 전구약물은, 예를 들어, 제WO2012/015760호에 기재되어 있다.
또한, ROCK 억제제는 ROCK1 및/또는 ROCK2 억제제인 것으로 또한 구상된다. 예시적인 ROCK1 억제제는 본원에 기재된 바와 같은 파수딜이다. 예시적인 ROCK1 및 ROCK2 억제제는 RhoXIII이다. 예시적인 ROCK2 억제제는 SLx-2119(KD025로도 알려짐)이다. ROCK 억제제는 표 1에 열거된 바와 같은 임의의 억제제 또는 이들의 조합물일 수 있다.
따라서, ROCK의 예시적인 억제제는 파수딜, Y27632(CAS: 146986-50-7), 하이드록시파수딜, H-1152-P(디메틸파수딜), Y27632, Y30141, Y32885(Wf536), Y39983(CAS: 203911-26-6), DW1865, SLx-2119(CAS: 911417-87-3), SR8046, SR6246, 리파수딜(Ripasudil), AS1892892, AR12141, AR12432, INS-117548, INS-115644, AT13148, RKI1447, SAR407899, 네타르수딜(Netarsudil)(AR-13324로도 알려짐), AR12286, PG286*, PG324**, ATS907, AMA0076, 티아조비빈(Thiazovivin), 아자벤즈이미다졸-아미노푸라잔스, H-0104 디하이드로클로라이드(CAS: 913636-88-1), DE-104, 올레핀, 이소퀴놀린(CAS: 119-65-3), 인다졸(CAS: 271-44-3), 피리딘알켄 유도체, H-1152 디클로라이드(CAS: 871543-07-6), XD-4000, HMN-1152, 4-(1-아미노알킬)-N-(4-피리딜)사이클로헥산-카르복사미드(경구 제형), 로스타틴, BA-210, BA-207, BA-215, BA-285, BA-1037, Ki-23095, VAS-012, 퀴나졸린, AR13154, AMA0428 및/또는 Rho XIII 또는 이들의 조합물을 포함하나, 이에 제한되지 않는다.
ROCK 억제제는 또한 파수딜, 하이드록시파수딜, 디메틸파수딜, Y27632, Y30141, Y32885, Y39983, DW1865, SLx-2119, SR8046, SR6246, 리파수딜, AS1892892, AR12141, AR12432, INS-117548, INS-115644, AT13148, RKI1447, SAR407899, 네타르수딜, AR12286, ATS907, AMA0076, 티아조비빈, H-0104 디하이드로클로라이드, 올레핀, 이소퀴놀린, 인다졸, H-1152 디클로라이드, 4-(1-아미노알킬)-N-(4-피리딜)사이클로헥산-카르복사미드, 퀴나졸린, Rho XIII, AR13154, AMA0428 및/또는 이들의 조합물로부터 선택될 수 있다.
ROCK 억제제는 또한 파수딜, Y27632, Y39983, SLx-2119, 리파수딜, INS-117548, INS-115644, AT13148, SAR407899, 네타르수딜, AR12286, ATS907, Rho XIII 및/또는 이들의 조합물로부터 선택될 수 있다.
본원에 기재된 ROCK 억제제는 하기 표에 추가로 특징화된다.
ROCK 억제제는 또한 파수딜, Y27632, 하이드록시파수딜, Y39983, SLx-2119, 리파수딜, ATS907, INS-117548, AT13148, SAR407899, 네타르수딜, AR12286, 또는 이들의 조합물로부터 선택될 수 있다.
ROCK 억제제는 또한 파수딜, Y27632, 하이드록시파수딜, Y39983, SLx-2119, 리파수딜, INS-117548, INS-115644, ATS907, AT13148, SAR407899, 네타르수딜, AR12286, PG286*, PG324**, ATS907, AMA0076 및 DE-104 또는 이들의 조합물로부터 선택될 수 있다.
ROCK 억제제는 또한 제WO 2009/158587호 및/또는 문헌[Feng et al. (2015) "Rho Kinase (ROCK) Inhibitors and Their Therapeutic Potential` including the information about clinical trials" J. Med. Chem. 2016, 59, 6, 2269-2300]에 개시된 바와 같은 ROCK 억제제일 수 있다.
추가로, ROCK 억제제는 본원에 기재된 임의의 적합한 ROCK 억제제의 염일 수 있다.
따라서, ROCK 억제제는 파수딜 HCl(이소퀴놀린 5-[(헥사하이드로-1H-1,4-디아제핀-1-일)설포닐]-N-(5-이소퀴놀린설포닐)-1,4-퍼-하이드로디아제핀; CAS No. 103745-39-7)과 같은 파수딜 또는 이의 염일 수 있다. 파수딜 HCl은 하기 화학식 I을 갖는다:
파수딜은 Rho 키나제, 특히 ROCK2의 강력하고 선택적인 억제제이다(Ono-Saito N, et al. (1999) "H-series protein kinase inhibitors and potential clinical applications" Pharmacol Ther, 82(2-3), 123-131).
ROCK 억제제는 또한 Rho XIII (1-(3-하이드록시벤질)-3-(4-(피리딘-4-일)티아졸-2-일)우레아; CAS: 1342278-01-6)과 같은 Rho-키나제 1과 Rho-키나제 2(ROCK1 및 ROCK2) 둘 모두의 억제제일 수 있다.
Rho XIII는 화학식 II를 갖는다: 
따라서, ROCK 억제제는 또한 파수딜 및/또는 RhoXIII일 수 있다.
본 발명에 따른 ROCK 억제제는 폐부종의 치료 또는 예방 방법에 사용될 수 있다. 이와 같이, 용어 "치료하는" 또는 "치료"는 증상 호전 또는 개선의 목적으로 바이러스 감염과 관련된 폐부종을 앓고 있는, 본원의 다른 곳에 정의된, 대상체에게 ROCK 억제제를 바람직하게는 의약 형태로 투여하는 것을 포함한다.
또한, ROCK 억제제는 ROCK 억제제의 투여 전의 정단 NKA 국소화와 비교하여 폐 상피 세포에서 바이러스(아르티큘라바이러스목, 모노네가바이러스 및/또는 분야바이러스목) 유도 정단 NKA 국소화를 감소시킴으로써 작용할 수 있다.
본원에서 사용되는 용어 "감소시키는"은 바이러스 유도 정단 NKA 국소화의 양이 ROCK 억제제의 투여 전에 존재하는 NKA 국소화와 비교하여 본원에 기재된 바와 같은 ROCK 억제제의 존재 하에 10%, 20%, 30%, 40%, 50%, 60%, 70%, 80%, 90%, 95%까지 또는 100%까지 감소된다는 것을 의미한다.
다시 말해서, ROCK 억제제는 ROCK 억제제의 부재 또는 ROCK 억제제의 투여 전과 비교하여 ROCK 억제제의 존재 하에 적어도 약 10%의 바이러스 감염(아르티큘라바이러스목, 모노네가바이러스 및/또는 분야바이러스목)의 결과로서 정단으로 이동하는 총 정단 NKA의 양을 감소시킨다.
본원에서 사용되는 용어 "예방하다", "예방" 또는 "예방법", 및 "예방하는"은 바이러스 감염과 관련된 폐부종을 얻거나 발병할 위험의 감소를 지칭한다. 또한 "예방"은 바이러스 감염과 관련된 폐부종의 재발의 감소 또는 억제를 의미한다. 특히, ROCK 억제제는 NKA의 기저측 국소화를 안정화시킴으로써 폐부종을 예방할 수 있다. 바람직하게는, "예방하다"는 총 NKA의 60%, 70%, 80%, 90%, 95% 또는 100%가 기저측으로 국소화됨을 의미한다. NKA의 국소화는 특히, 예를 들어, 실시예에 기재된 바와 같이 NKA에 대한 면역염색에 의해 결정될 수 있다.
본원에 정의된 바와 같은 ROCK 억제제는 폐부종을 치료 또는 예방하는 데 사용된다. 용어 "폐부종"은 당업자에게 공지되어 있으며, 특히 문헌[Matthay et al. (2019) "Acute Respiratiory distress Syndrome" Nat Rev Dis Primers, 5(1): 18, p. 1-52 as well as Mutlu et al. (2005) "mechanisms of pulmonary edema clearance" Am J Physiol Lung Cell Mol Physiol. 289: L685-695 and Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914]에 기재되어 있다. 이와 같이 "폐부종"은 폐의 조직 및 공기 공간, 특히 폐의 미세 기낭인 폐포에서의 체액 축적을 지칭한다.
따라서, 본원에서 사용되는 폐부종은 혈관외 폐수(EVLW)의 축적에 의해 특징화된다. 폐부종을 초래할 수 있는 한 가지 요인은 폐 모세관 투과성의 증가이다. 폐에서의 체액 축적, 즉, 폐부종은 바이러스 감염과 관련된 폐 손상으로 인한 것일 수 있다. 폐에서 생성된 체액 축적은 가스 교환을 손상시키고 호흡 곤란 또는 심지어 기계적 인공호흡 요구로 이어질 수 있다(Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914).
예를 들어, 종종 인플루엔자 바이러스 감염은 폐렴 및 소위 "급성 호흡 곤란 증후군"(ARDS)과 관련이 있다. 이는 특히 문헌[
amendys-Silva et al. (2011) "Acute respiratory distress syndrome caused by influenza B virus infection in a patient with diffuse large B-cell lymphoma" Case Rep Med. 2011; 2011: 647528]에 기재되어 있다. ARDS는, 전형적으로 문헌[Matthay et al. (2019) "Acute Respiratory Distress Syndrome" Nat rev Dis primers, 5(1):18]에 또한 기재된 바와 같은 호흡기 상피의 광범위한 손상에 의해 유발되는, 폐에서의 체액 축적의 한 형태이다.
폐부종은 흉부 엑스레이 또는 컴퓨터 단층촬영에 의해 진단될 수 있다. 흉부 엑스레이는 당업자에게 공지되어 있다. 이는 흉부에 영향을 미치는 상태를 진단하는 데 사용되는 흉부의 투영 방사선사진이다. 단순 흉부 방사선사진에서 폐부종을 광범위하게 평가하는 데 유용한 특징은 중심 폐정맥 울혈, 상엽 폐정맥 전환/폐정맥 울혈/수사슴 뿔 징후, 증가된 흉부 비율/심장 음영 크기를 포함하고; 근본적인 심장인성 원인 또는 연관성에 대한 평가에 유용하다. 폐 간질 부종의 특징은 다음을 포함할 수 있다: 기관지주위 둘러쌈 및 폐문주위 음영(perihilar haze), 중격선/컬리선, 엽간 균열의 비후. 폐의 폐포 부종의 특징: 배트윙 분포에서 전형적으로 공기 공간 혼탁은 공기 기관지조영술 흉막 삼출 및 엽간 균열의 체액('사라지는' 폐 가성종양 포함)을 가질 수 있다. 폐 모세관 쐐기 압력(PCWP)이 증가함에 따라 발생하는 단순 방사선사진 상의 징후의 일반적인 진행이 있다. 이러한 특징의 전부 또는 일부만이 단순 흉부 방사선사진에서 인지될 수 있는지 여부는 특정 병인에 따라 다르다. 또한, 폐부종은 일반적으로 양측 과정이지만, 특정 상황 및 병리학에서는 드물게 일방적인 것으로 나타날 수 있다.
폐 모세관 쐐기 압력(PCWP)은 팽창된 벌룬을 갖는 폐 카테터를 작은 폐 동맥 분지에 쐐기로 고정시킴으로써 측정된 압력이다. 이는 좌심방 압력을 추정한다. PCWP를 측정하는 방법은 특히 문헌[McIntyre et al. (1992) "A noninvasive method of predicting pulmonary-capillary wedge pressure" N Engl J Med. 327(24): 1715-20]에 기재되어 있다.
추가로, 폐부종은 등급 1일 수 있는 것으로 구상된다. 등급 1에서 폐 혈관 울혈은 흉부 방사선사진에서 검출될 수 있고, 구체적으로 페디칼 폭은 53 cm 초과로 검출될 수 있고/있거나 PCWP는 12-17 mmHg일 수 있다(Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914).
추가로, 폐부종은 등급 2일 수 있는 것으로 고려된다. 등급 2에서 간질 부종이 흉부 방사선사진에서 검출될 수 있고, 구체적으로 컬리 B 선 및/또는 기관지주위 둘러쌈(기관지 벽 상의 단부 비후)이 검출될 수 있고/있거나 PCWP는 17-25 mmHg일 수 있다(Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914).
추가로, 폐부종은 등급 3일 수 있는 것으로 구상된다. 등급 3에서, 폐포 부종의 증거는 흉부 방사선사진에서 검출될 수 있고, 구체적으로 폐 통합이 존재하는 것으로 검출될 수 있고/있거나 PCWP는 >25 mmHg일 수 있다.
따라서, 폐부종은 등급 1, 2 또는 3일 수 있다. 폐부종은 등급 1일 수 있다. 또한, 폐부종은 등급 2일 수 있는 것으로 구상된다. 또한, 폐부종은 등급 3일 수 있는 것으로 고려된다(Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914).
폐부종은 또한 특히 문헌[Assaad et al. (2018) "Assessment of Pulmonary Edema: Principles and Practice." Journal of Cardiothoracic and Vascular Anesthesia, 32(2), 901-914]에 기재된 바와 같이 폐 초음파 및/또는 경폐 열희석에 의해 검출될 수 있다.
본 발명은 ROCK 억제제가 본원에 개시된 바와 같이 바이러스 감염 시 폐 상피 세포에서 정단 NKA 국소화를 방지/감소시키는 것을 요구할 수 있다.
본원에서 사용되는 NKA는 임의의 적합한 NKA를 지칭한다. NKA는 당업자에게 공지되어 있으며, 특히 문헌[Mutlu et al. (2005) "mechanisms of pulmonary edema clearance" Am J Physiol Cell Mol phsiol 289: L685-L695]에 기재되어 있다. 이는 α- 및 β-서브유닛으로 구성된 이종이량체 단백질이다. α-서브유닛은 고에너지 포스페이트 결합을 절단하고 세포내 Na+를 세포외 K+로 교환하기 위한 촉매 부위를 갖는다. β-서브유닛은 이종이량체 조립 및 원형질막으로의 삽입을 제어하는 것으로 보이는 더 작은 글리코실화된 막관통 단백질이다. 둘 모두의 서브유닛은 기능성 NKA에 필요하다. 따라서, NKA는 α 및 β 서브유닛을 포함한다.
예를 들어, NKA는 SEQ ID NO. 4-7의 서열 또는 SEQ ID NO. 4-7의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열 중 어느 하나의 α 서브유닛을 포함할 수 있다.
추가로 또는 대안적으로, NKA는 SEQ ID NO. 1-3의 서열 또는 SEQ ID NO. 1-3의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열 중 어느 하나의 β 서브유닛을 포함할 수 있다.
따라서, NKA는 SEQ ID NO. 4-7의 서열 또는 SEQ ID NO. 4-7의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열 중 어느 하나의 α 서브유닛 및 SEQ ID NO. 1-3의 서열 또는 SEQ ID NO. 1-3의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열 중 어느 하나의 β 서브유닛을 포함할 수 있다.
본 발명에 따르면, SEQ ID NO. 1-15와 같은 2개 이상의 폴리펩타이드 서열의 맥락에서 "동일한" 또는 "동일성 퍼센트"라는 용어는, 비교 범위에 걸쳐, 또는 당 분야에 공지된 서열 비교 알고리즘을 사용하여, 또는 수동 정렬 및 육안 검사에 의해 측정된 바와 같이 지정된 영역에 걸쳐 최대 일칭성에 대해 비교하고 정렬하는 경우, 동일하거나, 동일한 아미노산의 특정 백분율(예를 들어, 적어도 85%, 90%, 95%, 96%, 97%, 98% 또는 99% 동일성)을 갖는 둘 이상의 서열 또는 하위서열을 지칭한다. 예를 들어, 80% 내지 95% 이상의 서열 동일성을 갖는 서열은 실질적으로 동일한 것으로 간주된다. 이러한 정의는 또한 시험 서열의 상보체에도 적용된다. 당업자는 예를 들어, 당 분야에 공지된 바와 같은 CLUSTALW 컴퓨터 프로그램(Thompson Nucl. Acids Res. 2 (1994), 4673-4680) 또는 FASTDB(Brutlag Comp. App. Biosci. 6 (1990), 237-245)를 기반으로 한 것들과 같은 알고리즘을 이용하여 서열 사이/서열 간 동일성 퍼센트를 결정하는 방법을 알 것이다.
또한 BLAST 및 BLAST 2.6 알고리즘(Altschul Nucl. Acids Res. 25 (1977), 3389-3402)이 당업자에게 이용 가능하다. 아미노산 서열에 대한 BLASTP 프로그램은 기본적으로 6의 단어 크기(W), 10의 예상 임계값, 및 둘 모두의 가닥의 비교를 디폴트로서 사용한다. 또한, BLOSUM62 스코어링 매트릭스(Henikoff Proc. Natl. Acad. Sci., USA, 89, (1989), 10915; Henikoff and Henikoff (1992) 'Amino acid substitution matrices from protein blocks.' Proc Natl Acad Sci U S A. 1992 Nov 15;89(22):10915-9)가 사용될 수 있다.
예를 들어, 기본적인 국소 정렬 검색 도구(Basic Local Alignment Search Tool)를 나타내는 BLAST2.6(Altschul, Nucl. Acids Res. 25 (1997), 3389-3402; Altschul, J. Mol. Evol. 36 (1993), 290-300; Altschul, J. Mol. Biol. 215 (1990), 403-410)을 이용하여 국소 서열 정렬을 검색할 수 있다.
NKA는, 예를 들어, 건강한 대상체에서 폐 상피 세포의 기저측 표면에 위치한다. NKA는 ATP를 소비함으로써 이온을 수송한다. 구체적으로, 이는 칼륨 유입과 교환하여 세포 밖으로 Na+ 이온을 펌핑한다. 이러한 방식으로, 이는 원형질막에 걸쳐 Na+ 및 칼륨 농도구배를 유지한다.
본원에서 사용되는 표현 "폐 상피 세포"는 임의의 폐 상피 세포를 의미한다. 당업자는, 특히 문헌[Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730] 및 문헌[Crystal et al. (2008) "Airway epithelial cells" Proc Am Thorac Soc, vol. 5, pp. 772-777]에 기재되어 있는, 폐의 상이한 상피 세포를 알고 있다. 일반적으로, 폐 상피 세포는 분극화된다. 이는 폐 상피 세포가 정단 및 기저 쪽을 갖는다는 것을 의미한다.
폐 상피 세포는 기관 상피 세포, 기관지 상피 세포 또는 폐포 상피 세포일 수 있다.
당업자는 세포가 이러한 상피 세포 중 하나에 속하는지를 결정할 수 있다. 예를 들어, 당업자는 발현 코넥신 43을 검출하기 위해 면역조직화학을 수행할 수 있다. 코넥신 43은 폐포 상피 세포(AT1 및 AT2 세포 둘 모두)에서, 뿐만 아니라 기관 상피 세포 및 기관지 상피 세포 상에서도 발현된다(Johnson and Koval (2009) "Cross-Talk Between Pulmonary Injury, Oxidant Stress, and Gap Junctional Communication." Antioxidans and Redox Signaling, vol. 11, number 2, pp. 356-367). 코넥신 43은 SEQ ID NO. 8에 나타나 있는 바와 같은 서열을 가질 수 있거나 SEQ ID NO. 8의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열일 수 있다.
따라서, 본원에 기재된 바와 같은 폐 상피 세포는 코넥신 43을 발현하는 세포일 수 있다. 폐 상피 세포는 SEQ ID NO. 8에 나타나 있는 바와 같은 코넥신 43을 발현하는 세포일 수 있거나 SEQ ID NO. 8의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열일 수 있다.
보다 구체적으로, "기관 상피 세포"는 약 20에서 25 분지(큰 기도; 기관 상피 세포)로 연장될 수 있다. 기관 상피 세포는 기관 및 더 큰 기관지를 라이닝한다.
"기관지 상피 세포"는 26에서 223 분지(작은 기도, 기관지 상피 세포)로 연장될 수 있다. 기관지 상피 세포는 세기관지 및 더 작은 기관지를 라이닝한다.
기관 상피 세포와 기관지 상피 세포 둘 모두는 소위 섬모 가중층화된 원주 상피에 국소화된다.
따라서, 섬모 가중층화된 원주 상피는 기관뿐만 아니라 상기도의 내층에서도 발견된다. 가중층화된 상피는 인간에서 기관으로부터 원위 세기관지로 연장될 수 있다. 예를 들어, 이는 20에서 25 분지(큰 기도; 기관 상피 세포)로 및 26에서 223 분지(작은 기도, 기관지 상피 세포)로 연장될 수 있다. 가중층화된 상피는 다중 층을 갖는 것으로 보이지만, 실제로는 단일 시트의 세포로만 구성된다. 개별 원주 세포 내의 핵의 위치결정은 이러한 착시를 야기한다. 이러한 핵은 다양한 수준으로 발견되어 계층화된 외관을 생성한다.
기관 및 기관지 상피의 주요 세포 유형은 섬모 세포, 분비 세포 및 기저 세포이다(Crystal et al. (2008) "Airway epithelial cells" Proc Am Thorac Soc, vol. 5, pp. 772-777; Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730).
결과적으로, 폐 상피 세포는 기관지 상피 세포일 수 있는 것으로 구상된다. 추가로, 폐 상피 세포는 폐포 상피 세포일 수 있는 것으로 구상된다. 기관지 또는 기관 상피 세포는 섬모 세포, 분비 세포 또는 기저 세포일 수 있다.
본원에서 사용되는 "섬모 세포"는 아래에 있는 기저판에 부착된 얇고 테이퍼링된 염기를 갖는다. 세포는 또한 대부분의 물질에 대해 물리적으로 불투과성인 장벽을 형성하고, 데스모솜에 의해 서로 및 기저 세포에 대해 측방향으로 장벽을 형성하는 밀착 접합부에 의해 이들의 정단 표면에서 서로 부착될 수 있다. 수많은 미세융모를 함유하는 세포간 공간이 세포 사이에 존재할 수 있다(Crystal et al. (2008) "Airway epithelial cells" Proc Am Thorac Soc, vol. 5, pp. 772-777; Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730).
분비 세포는 배상 세포, 클라라 세포, 내강 분비 세포 또는 신경내분비 세포일 수 있다.
본원에서 사용되는 "배상 세포"는 상부 및 하부 기도의 표면 상피에 위치하며, 기도를 코팅하고 제거될 미립자를 포획하기 위해 점액을 생성할 수 있다.
본원에서 사용되는 "클럽 세포"로도 지칭되는 "클라라 세포"(비섬모 세기관지 분비 세포)는 더 원위 기도를 라이닝하며, 따라서 이러한 세포 유형은 주로 막성 세기관지에서 발견될 수 있다. 이들의 형상은 종종 원주형이거나 (더 원위 기도에서) 입방형이다. 이들은 성숙한 계면활성제 단백질 A, B, D 및 여러 해독 효소를 분비할 수 있다.
본원에서 사용되는 "내강 분비 세포"는 비-섬모 세포이다. 내강 전구 세포는 휴식 조건 하에서 대부분의 증식 세포를 차지한다(Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730).
신경내분비 세포는 기저부에서 기저막에 부착될 수 있고, 기도 표면을 향해 연장되고 기도 표면에 도달할 수 있거나 도달하지 않을 수 있는 테이퍼링된 정단을 가질 수 있다. 세포의 주요 기능은 펩타이드의 분비이다(Crystal et al. (2008) "Airway epithelial cells" Proc Am Thorac Soc, vol. 5, pp. 772-777; Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730).
본원에서 사용되는 "기저 세포"는 기저막과 접촉하지만 기도 내강과 접촉하지 않을 수 있는 비교적 부유한 세포 유형이다. 기저 세포는 표면 상피 아래에 위치할 수 있고 섬모 세포와 분비 세포 둘 모두의 전구체로서 작용할 수 있다. 이들은 손상 후 기도 상피의 재생에 중요한 역할을 한다. 중간 필라멘트 단백질의 특징적인 서브세트(케라틴 5 [K5], K6, K14, 및 K16)의 발현은 기저 세포를 내강 상피 세포와 구별할 수 있다(Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730).
따라서, 본원에서 사용되는 폐 상피 세포는 섬모 또는 비-섬모 기관 상피 세포일 수 있다. 또한, 본원에서 사용되는 폐 상피 세포는 섬모 또는 비-섬모 기관지 상피 세포일 수 있는 것으로 고려된다.
섬모 기관 또는 기관지 세포는 본원에 기재된 바와 같은 섬모 세포일 수 있다. 비-섬모 기관/기관지 세포는 신경내분비 세포, 내강 분비 세포, 클라라 세포 또는 기저 세포일 수 있다.
상피 세포는 섬모 가중층화된 원주 상피 또는 단순 편평 상피에 국소화될 수 있다. 본원에 기재된 바와 같이, 기관 및 기관지 상피 세포는 용어 "섬모 가중층화된 원주 상피"에 포함된다.
다른 한 편으로, 본원에서 폐포 상피로도 지칭되는 단순 편평 상피는 223 분지(폐포) 후에 존재하며, 타입 I 및 타입 II 세포를 포함한다. 따라서, 폐포 상피는 타입 I 및 II 폐포 상피, 또는 타입 I(AT1) 및 타입 II(AT2) 폐포 상피 세포의 혼합을 포함한다. 폐포는 호흡기에서 가장 작은 기능 단위이며, 폐에서 모세관과 산소 및 이산화탄소와 같은 가스 교환을 담당한다. 폐포 상피 단층은 얇고, 편평 타입 I 세포(가스 교환을 허용하는 AT1) 및 입방형 타입 2 세포(AT2, 폐 확장을 가능하게 하는 계면활성제를 생산함)로 이루어진다. 둘 모두의 세포는 폐포로부터 이온 및 체액을 수송하여 건조한 공기 공간을 유지한다.
AT1 세포는 폐의 내부 표면적의 약 95%를 덮는다. 이들은 종종 인접한 폐포로 연장될 수 있는 다수의 정단 표면을 갖는 분지된 세포이다. AT1 세포의 정단 표면적은 대부분의 세포와 비교하여 크지만(즉, 인간 AT1 세포의 경우 약 5,000 μm2), 이들은 매우 얇은 세포(즉, 0.2 μm 깊이)이다. 가스 교환 장벽은 융합된 기저막에 의해 접합된 AT1 및 내피 세포로 구성된다.
세포가 타입 I 세포인지 결정하기 위한 마커는 당업자에게 공지되어 있으며, 특히 문헌[McElroy and Kasper (2004) "The use of alveolar epithelial type I cell-selective markers to investigate lung injury and repair" European Respiratory Journal 24: 664-673]에 기재되어 있다. 예를 들어, 세포가 타입 I 세포인지 결정하기 위해, 당업자는 면역조직화학을 수행하여 RTI40/TIα 단백질, HTI56 및/또는 Na+/K+-ATPase α2-이소형(α2-이소형은 SEQ ID NO. 5에 도시됨) 중 하나 이상의 발현을 검출할 수 있다. Na+/K+-ATPase α2-이소형은 SEQ ID NO. 5에 나타나 있는 바와 같은 서열을 가질 수 있거나 SEQ ID NO. 5의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열일 수 있다.
추가로 또는 대안적으로, 당업자는 특히 문헌[Marconett et al. (2016) "Cross-Species Transcriptome Profiling Identifies New Alveolar Epithelial Type I Cell-Specific Genes" AJRCMB, vol 56, no. 3]에 기재된 바와 같이 면역조직화학을 수행하여 진행성 글리코실화 최종 산물-특이적 수용체(AGER, 이전에 RAGE), 포도플라닌(PDPN, 이전에 T1a), 카베올린1(CAV1), HOPX, GRAM 도메인 2(GRAMD2) 중 하나 이상의 발현을 검출할 수 있다. 인간 GRAMD2 이소형 A 및 B는 SEQ ID NO. 9 및 10에 나타나 있다. GRAMD2는 SEQ ID NO. 9 또는 10에 나타나 있는 서열을 가질 수 있거나 SEQ ID NO. 9 또는 10의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열일 수 있다.
따라서, 본원에 기재된 바와 같은 폐 상피 세포는 Na+/K+-ATPase α2-이소형 및/또는 GRAMD2를 발현하는 세포일 수 있다. 폐 상피 세포는 SEQ ID NO. 5 또는 SEQ ID NO. 5의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열에 나타나 있는 바와 같은 Na+/K+-ATPase α2-이소형 및/또는 SEQ ID NO. 9 또는 10 또는 SEQ ID NO. 9 또는 10의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열에 나타나 있는 바와 같은 GRAMD2를 발현하는 세포일 수 있다.
AT2는 AT1 세포 사이에 위치한 입방형 세포이고, 특징적인 층상체 및 정단 미세융모를 함유한다. AT2 세포는 폐 계면활성제의 생산, 분비 및 재흡수 및 숙주 방어에 중요한 면역조절 단백질의 합성 및 분비를 포함하는 다수의 공지된 기능을 갖는다. 세포가 타입 II 세포인지 결정하기 위한 마커는 당업자에게 공지되어 있으며, 특히 문헌[McElroy and Kasper (2004) "The use of alveolar epithelial type I cell-selective markers to investigate lung injury and repair" European Respiratory Journal 24: 664-673]에 기재되어 있다. 예를 들어, 세포가 타입 II 세포인지 결정하기 위해, 당업자는 면역조직화학을 수행하여 폐 계면활성제-관련 단백질 C(SP-C)의 발현을 검출할 수 있다. SP-C은 SEQ ID NO. 11에 나타나 있는 바와 같은 서열을 가질 수 있거나 SEQ ID NO. 11의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열일 수 있다.
따라서, 본원에 기재된 바와 같은 폐 상피 세포는 SP-C를 발현하는 세포일 수 있다. 폐 상피 세포는 SEQ ID NO. 11 또는 SEQ ID NO. 11의 서열에 대해 70%, 80%, 90%, 95%, 99% 서열 동일성을 갖는 서열에 나타나 있는 바와 같은 SP-C을 발현하는 세포일 수 있다.
따라서, 본원에 기재된 바와 같은 폐 상피 세포는 또한 SP-C를 발현하는 세포 및 Na+/K+-ATPase α2-이소형 및/또는 GRAMD2를 발현하는 세포를 포함할 수 있다.
유형 I 및 유형 II 세포 둘 모두는 당업자에게 공지되어 있으며, 특히 문헌[Rackley et al. (2012) "Building and maintaining the epithelium of the lung" The Journal of Clinical investigation, vol. 122, no. 8, pp. 2724-2730, Crystal et al. (2008) "Airway epithelial cells" Proc Am Thorac Soc, vol. 5, pp. 772-777] 및 문헌[McElroy and Kasper (2004) "The use of alveolar epithelial type I cell-selective markers to investigate lung injury and repair" European Respiratory Journal 24: 664-673]에 기재되어 있다. 특히, 본원에 기재된 바와 같은 NKA는 AT1 및 AT2 세포 둘 모두에서 발현된다.
따라서, 폐 상피 세포는 폐포 상피 세포일 수 있다. 폐 상피 세포는 또한 타입 I(AT1) 및/또는 타입 II(AT2) 세포일 수 있다.
본원에 개시된 바와 같은 폐 상피 세포는 또한 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스에 감염된 폐 상피 세포, 바람직하게는 폐포 상피 세포일 수 있다. 추가로, 오르토믹소바이러스과(아르티큘라바이러스목), 아레나바이러스과(Arenaviridae), 한타바이러스과(Hantaviridae), 미포바이러스과(Mypoviridae), 나이로바이러스과(Nairovirdae), 페리분야바이러스과(Peribunyaviridae), 페뉴바이러스과(Phenuviridae)(분야바이러스목), 보르나바이러스과(Bornaviridae), 필로바이러스과(Filoviridae), 파라믹소바이러스과(Paramyxoviridae) 및/또는 순바이러스과(Sunviridae)(모노네가바이러스목)의 바이러스에 감염된 폐 상피 세포, 바람직하게는 폐포 상피 세포가 구상된다.
또한, 알파인플루엔자바이러스(Alphainfluenzavirus), 베타인플루엔자바이러스(Betainfluenzavirus), 델타인플루엔자바이러스(Deltainfluenzavirus), 감마인플루엔자바이러스(Gammainfluenzavirus) 속, 바람직하게는 알파인플루엔자바이러스(아르티큘라바이러스목; 오르토믹소바이러스과), 맘만타바이러스과(Mammantavirinae) 아과, 바람직하게는 로안 바이러스(Loan virus), 모바트 바이러스(Mobat virus), 오르토한타바이러스(Orthohantavirus) 또는 토팀바이러스(Thottimvirus) 속(분야바이러스목; 한타바이러스과), 및/또는 뉴모바이러스과 속(모노네가바이러스목, 파라마이옥소바이러스과)의 바이러스 감염된 폐 상피 세포, 바람직하게는 폐포 상피 세포가 고려된다.
추가로, H1N1-, H1N2-, H2N2-, H3N2-, H5N1-, H6N1-, H7N2-, H7N3-, H7N7-, H7N9, H9N2-, H10N7-, H10N8- 또는 H5N1-서브타입(아르티큘라바이러스목; 오르토믹소바이러스과, 알파인플루엔자바이러스 속), 퓨말라 바이러스(Puumala virus), 신 놈브레 바이러스(Sin Nombre virus), 서울 바이러스(Seoul virus), 한탄 바이러스(Hantaan virus), 도브라바-벨그라드 바이러스(Dobrava-Belgrad virus), 사레마 바이러스(Saaremaa virus), 포 코너즈 바이러스(Four corners virus) 또는 안데스 바이러스(Andes virus)(분야바이러스목; 한타바이러스과, 맘만타바이러스과 아과, 오르토한타바이러스 속) 또는 메타뉴모바이러스(Metapneumovirus), 오르토뉴모바이러스(Orthopneumovirus), 예컨대, 인간 오르토뉴모바이러스(Human orthopneumovirus), 예컨대, 인간 호흡기 세포융합 바이러스 A(Human respiratory syncytial virus A), 인간 호흡기 세포융합 바이러스 B(Human respiratory syncytial virus B) 또는 미분류 인간 호흡기 세포융합 바이러스, 개 뉴모바이러스(Canine pneumovirus), 고양이 뉴모바이러스(Feline pneumovirus), 양 호흡기 세포융합 바이러스(Ovine respiratory syncytial virus), 양 호흡기 세포융합 바이러스(Ovine respiratory syncytial virus)(균주 WSU 83-1578), 뉴모바이러스(Pneumovirus), 호흡기 세포융합 바이러스(Respiratory syncytial virus), 돼지 뉴모바이러스(Swine pneumovirus) 또는 뉴모바이러스(Pneumovirus) 종(모노네가바이러스목, 파라마이옥소바이러스과, 뉴모바이러스과 속)에 감염된 폐 상피 세포, 바람직하게는 폐포 상피 세포가 구상된다.
또한, 폐 상피 세포, 바람직하게는 폐포 상피 세포는 본원에 개시된 바와 같은 2종 이상의 상이한 바이러스에 감염되는 것으로 고려된다.
당업자는 폐 상피 세포가 본원에 개시된 바와 같은 바이러스에 감염되었는지를 검출하는 방법을 알고 있다. 예를 들어, 당업자는 폐 상피 세포 조직의 조직 샘플을 취할 수 있다. 이후, 당업자는 관심 바이러스의 바이러스 게놈 서열 또는 서열 단편이 이러한 세포에 존재하는지 분석하기 위해 이러한 샘플에 대해 PCR을 수행할 수 있다.
일반적으로 폐 상피 세포에 대한 모델은 실시예에서 사용되는 바와 같은 및 특히 문헌[Peteranderl et al. (2019) "Influenza A virus infection induces apical redistribution of Na+, K+-ATPase in lung epithelial cells in vitro and in vivo" American Journal of Respiratory Cell and Molecular Biology Volume 61 Number 3, pp. 395-397]에 기재된 바와 같은 Calu3 세포주(American Type Culture Collection(매너서스, VA, USA)로부터 입수할 수 있는 인간 선암종 기관지 상피 세포)이다.
본원에서 사용되는 ROCK 억제제는 폐 상피 세포에서 정단 NKA 국소화를 방지할 수 있다. 본원에 개시된 바와 같이, NKA는 건강한 대상체에서 분극화된 폐 상피 세포의 원형질막 내에 기저측으로 국소화되거나 이에 회합된다. 기저측 국소화는 또한 손상 동안 유지될 수 있다.
특히, 상피 세포는 조밀한 접합부, 데스모솜 및 부착물 접합부를 통해 서로 부착되어, 동물/인간의 신체 및 내부 강(예를 들어, 호흡기관, 소화관 및 순환계)의 표면을 라이닝하는 세포 시트를 형성한다. 이러한 세포는 신체의 외부 표면, 또는 내부 공동의 내강을 향하는 정단 막, 및 내강으로부터 멀리 배향된 기저측 막에 의해 정의되는 정단-기저 극성을 갖는다. 기저측 막은 세포-세포 접합부가 이웃 세포를 연결하는 측부 막 및 세포가 기저막에 부착되는 기저막 둘 모두를 지칭하며, 세포외 기질 단백질의 얇은 시트가 상피 시트를 하부 세포 및 결합 조직으로부터 분리하고 있다(Wu and Mlodzik (2009). "A quest for the mechanism regulating global planar cell polarity of tissues". Trends in Cell Biology. 19 (7): 295-305).
따라서, 용어 "기저측"은 세포막으로 지칭될 때, 인접한 세포 및 밑에 있는 결합 조직을 향하는 원형질막의 분획이다. 유사하게, 세포막을 지칭하는 용어 "정단"은 공동의 내강을 향하는 세포막의 분획을 의미하는 것이다.
도 1은 NKA 국소화에 대한 ROCK 억제제의 효과를 보여준다. 건강한 세포에서 NKA는 기저측으로 국소화된다. 바이러스 감염 시 NKA는 기저측으로 국소화되지만 또한 정단에서도 국소화된다. 인플루엔자 A 바이러스에 감염된 Calu 3 세포의 막에 대한 NKA 분포는, 예를 들어, 광학적 가시화 방법에 의해 검출될 수 있다. 예를 들어, NKA는 실시예 7에 기재된 바와 같이 항체 염색 및 공초점 레이저-스캐닝 현미경에 의해 가시화될 수 있다.
본 발명은 또한 ROCK 억제제가 ROCK 억제제의 투여 전 정단 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킬 수 있는 것을 구상한다.
본 발명자들은 놀랍게도 바이러스 감염이 폐 상피 세포에서 정상적으로 존재하는 기저측 NKA 국소화에 추가하여 폐 상피 세포에서 정단 NKA 국소화를 초래한다는 것을 발견하였다. ROCK 억제제의 투여 시, 폐 상피 세포에서 NKA 국소화의 기저측 국소화는 대부분 회복될 수 있는 반면, 폐 상피 세포에서 NKA 국소화의 정단 국소화는 ROCK 억제제를 투여하기 전의 상황과 비교하여 감소된다.
따라서, ROCK 억제제는 ROCK 억제제의 투여 전 존재하는 폐 상피 세포에서 정단으로 국소화된 NKA와 비교하여 폐 상피 세포에서 바이러스-유도 정단 국소화된 NKA의 5%, 10%, 15%, 20%, 30%, 40%, 50% 60%, 70%, 80%, 90% 95%, 99% 또는 100% 감소를 야기한다.
본원에 개시된 바와 같은 폐부종은 바이러스 감염과 관련되고, 여기서 바이러스는 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목이다.
아르티큘라바이러스목의 바이러스는 바람직하게는 오르토믹소바이러스과이다.
오르토믹소바이러스과의 바이러스는 알파인플루엔자바이러스, 베타인플루엔자바이러스, 델타인플루엔자바이러스, 감마인플루엔자바이러스, 이사바이러스, 쿠아란자바이러스, 토고토바이러스, 미분류 오르토믹소바이러스과 속일 수 있다.
알파인플루엔자바이러스 속은 헤마글루티닌(HA) 항원성 서브타입(HX) 및 뉴라미니다제(NA) 항원성 서브타입(NY)의 임의의 조합을 보유하는 인플루엔자 A 바이러스일 수 있다. 예를 들어, HA는 서열 SEQ ID NO. 12 또는 13, 또는 SEQ ID NO. 11 또는 12의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열일 수 있다. 추가로 또는 대안적으로, NA는 서열 SEQ ID NO. 14 또는 15, 또는 SEQ ID NO. 14 또는 15의 서열에 대해 70%, 75%, 80%, 85%, 90%, 95%, 97%, 98%, 99% 서열 동일성을 갖는 서열일 수 있다.
알파인플루엔자바이러스의 비제한적인 예는 H1N1-, H1N2-, H2N2-, H3N2-, H5N1-, H6N1-, H7N2-, H7N3-, H7N7-, H7N9, H9N2-, H10N7-, H10N8- 또는 H5N1-서브타입을 포함한다. 일 구현예에서, 인플루엔자 A 바이러스는 H1N1-서브타입이다. 다른 구현예에서, 인플루엔자 A 바이러스는 H3N2-, H5N1- 및 H7N9-서브타입이다. 인플루엔자 A 바이러스는 또한 H3N2-, H5N1-, H1N1- 및 H7N9-서브타입일 수 있다. 인플루엔자 A 바이러스는 또한 균주 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)일 수 있다.
바이러스는 분야바이러스(Bunyavirales) 목일 수 있다. 당업자는 어떤 바이러스가 분야바이러스목에 속하는지 알고 있다. 모든 분야바이러스목은 세 부분으로 분할되는 음성-센스 RNA 게놈을 갖는다.
분야바이러스목의 바이러스는 아레나바이러스과, 한타바이러스과, 마이포바이러스과, 나이로바이러스과, 페리분야바이러스과, 또는 페뉴바이러스과일 수 있다.
아레나바이러스과의 바이러스는 맘마레나바이러스(Mammarenavirus) 속일 수 있다. 맘마레나바이러스 속의 비제한적인 예는 특히 입피-바이러스(Ippy-Virus; IPPYV), 라사-바이러스(Lassa-Virus; LASV), 루조-바이러스(Lujo-Virus; LUJV), 룬크-바이러스(Lunk-Virus; NKS-1), 림프루성 맥락수막염 바이러러스(Lymphocytic choriomeningitis virus; LCMV), 모발라-바이러스(Mobala-Virus; MOBV), 및 모페이다-바이러스(Mopeia-Virus; MOPV) 종을 포함한다.
분야바이러스목의 바이러스는 바람직하게는 한타바이러스과이다. 한타바이러스과의 바이러스는 맘만타바이러스과 아과일 수 있다. 맘만타바이러스과 아과의 바이러스는 로안 바이러스, 모바트 바이러스, 오르토한타바이러스 또는 토팀바이러스 속일 수 있다.
오르토한타바이러스 속은 특히 퓨말라 바이러스, 신 놈브레 바이러스, 서울 바이러스, 한탄 바이러스, 도브라바-벨그라드 바이러스, 사레마 바이러스, 포 코너즈 바이러스 및 안데스 바이러스를 포함한다.
나이로바이러스과의 바이러스는 오르토나이로바이러스 또는 스트리와바이러스 속일 수 있다. 오르토나이로바이러스는 크리미아-콩고 출혈열 오르토나이로바이러스(Crimean-Congo hemorrhagic fever orthonairovirus; CCHFV) 종일 수 있다.
페리분야바이러스과의 바이러스는 오르토분야바이러스 또는 파쿠바이러스 속일 수 있다.
오르토분야바이러스 속은 아이노 오르토분야바이러스(Aino orthobunyavirus), 아카바네 오르토분야바이러스(Akabane orthobunyavirus), 안헴비 오르토분야바이러스(Anhembi orthobunyavirus), 아노펠레스 B 오르토분야바이러스(Anopheles B orthobunyavirus), 바타이 오르토분야바이러스(Batai orthobunyavirus), 바타마 오르토분야바이러스(Batama orthobunyavirus), 베르티오가 오르토분야바이러스(Bertioga orthobunyavirus), 분야웨라 오르토분야바이러스(Bunyamwera orthobunyavirus), 부토윌로우 오르토분야바이러스(Buttonwillow orthobunyavirus), 브왐바 오르토분야바이러스(Bwamba orthobunyavirus), 카케 밸리 오르토분야바이러스(Cache Valley orthobunyavirus), 카코에이라 포르테이라 오르토분야바이러스(Cachoeira Porteira orthobunyavirus), 카핌 오르토분야바이러스(Capim orthobunyavirus), 카라파루 오르토분야바이러스(Caraparu orthobunyavirus), 카투 오르토분야바이러스(Catu orthobunyavirus), 포르트 쉐르만 오르토분야바이러스(Fort Sherman orthobunyavirus), 감보아 오르토분야바이러스(Gamboa orthobunyavirus), 구아마 오르토분야바이러스(Guama orthobunyavirus), 구아로아 오르토분야바이러스(Guaroa orthobunyavirus), 이아코 오르토분야바이러스(Iaco orthobunyavirus), 이레샤 오르토분야바이러스(Ilesha orthobunyavirus), 인그와부마 오르토분야바이러스(Ingwavuma orthobunyavirus), 자토발 오르토분야바이러스(Jatobal orthobunyavirus), 켄그 코이 오르토분야바이러스(Kaeng Khoi orthobunyavirus), 케이스톤 오르토분야바이러스(Keystone orthobunyavirus), 라 크로세 오르토분야바이러스(La Crosse orthobunyavirus), 마카우아 오르토분야바이러스(Macaua orthobunyavirus), 마드리드 오르토분야바이러스(Madrid orthobunyavirus), 마구아리 오르토분야바이러스(Maguari orthobunyavirus), 만자닐라 오르토분야바이러스(Manzanilla orthobunyavirus), 마리투바 오르토분야바이러스(Marituba orthobunyavirus), 메르메트 오르토분야바이러스(Mermet orthobunyavirus), 오리보카 오르토분야바이러스(Oriboca orthobunyavirus), 오로포우케 오르토분야바이러스(Oropouche orthobunyavirus), 파토이스 오르토분야바이러스(Patois orthobunyavirus), 페아톤 오르토분야바이러스(Peaton orthobunyavirus), 사보 오르토분야바이러스(Sabo orthobunyavirus), 산고 오르토분야바이러스(Sango orthobunyavirus), 사투페리 오르토분야바이러스(Sathuperi orthobunyavirus), 슈말렌베르크 오르토분야바이러스(Schmallenberg orthobunyavirus), 슈니 오르토분야바이러스(Shuni orthobunyavirus), 심부 오르토분야바이러스(Simbu orthobunyavirus), 스노우슈 하레 오르토분야바이러스(Snowshoe hare orthobunyavirus), 소로로카 오르토분야바이러스(Sororoca orthobunyavirus), 타히나 오르토분야바이러스(Tahyna orthobunyavirus), 예컨대, 타히나 바이러스(Tahyna virus; TAHV), 타타구인 오르토분야바이러스(Tataguine orthobunyavirus), 테테 오르토분야바이러스(Tete orthobunyavirus), 우탄가 오르토분야바이러스(Utinga orthobunyavirus), 올크베르크 오르토분야바이러스(Wolkberg orthobunyavirus), 웨오마이아 오르토분야바이러스(Wyeomyia orthobunyavirus), 또는 젤라 오르토분야바이러스(Zegla orthobunyavirus) 종을 포함할 수 있다. 오르토분야바이러스 속은 또한 바아칼 바이러스(Baakal virus; BKAV)를 포함할 수 있다.
파쿠바이러스 속은 파쿠이 바이러스(Pacui virus; PACV), 리오 프레토 다 에바 바이러스(Rio Preto da Eva virus; RPEV) 및 타피라페 바이러스(Tapirape virus; TPPV) 종을 포함할 수 있다.
페뉴이바이러스과의 바이러스는 반얀그바이러스(Banyangvirus), 고우코바이러스(Goukovirus), 또는 플레보바이러스(Phlebovirus) 속이다.
플레보바이러스 속은 틱-보르넨 플레보바이러스(tick-borne Phlebovirus) 종일 수 있다.
모네가티바이러스목의 바이러스는 보르나바이러스과, 필로바이러스과, 파라믹소바이러스과 또는 순바이러스과일 수 있다. 예를 들어, 모노네가티바이러스목의 바이러스는 뉴모바이러스과 속일 수 있다.
보르나바이러스과의 바이러스는 카르보바이러스, 오르토보르나바이러스 또는 미분류 보르나바이러스과 속일 수 있다.
필로바이러스과의 바이러스는 쿠에바바이러스, 에볼라바이러스 또는 마르부르그바이러스 속일 수 있다.
파라믹소바이러스과의 바이러스는 아부알바이러스과, 아불라바이러스, 오르토파라믹소바이러스과, 루불라바이러스과, 루불라바이러스, 미분류 파라믹소바이러스과, 메타뉴모바이러스, 오르토뉴모바이러스, 또는 뉴모바이러스과 속일 수 있다.
뉴모바이러스과 속의 바이러스는 메타뉴모바이러스, 오르토뉴모바이러스, 예컨대, 인간 오르토뉴모바이러스, 예컨대, 인간 호흡기 세포융합 바이러스 A, 인간 호흡기 세포융합 바이러스 B 또는 미분류 인간 호흡기 세포융합 바이러스, 개 뉴모바이러스, 고양이 뉴모바이러스, 양 호흡기 세포융합 바이러스, 양 호흡기 세포융합 바이러스(균주 WSU 83-1578), 뉴모바이러스, 호흡기 세포융합 바이러스, 돼지 뉴모바이러스 또는 뉴모바이러스 종일 수 있다.
따라서, 본원에 개시된 바이러스는 오르토믹소바이러스과(아르티큘라바이러스목), 아레나바이러스과, 한타바이러스과, 마이포바이러스과, 나이로바이러스과, 페리분야바이러스과, 페뉴바이러스과(분야바이러스목), 보르나바이러스과, 필로바이러스과, 파라믹소바이러스과 또는 순바이러스과(모노네가바이러스목)의 바이러스일 수 있다.
따라서, 본원에 개시된 바이러스는 알파인플루엔자바이러스, 베타인플루엔자바이러스, 델타인플루엔자바이러스, 감마인플루엔자바이러스, 바람직하게는 알파인플루엔자바이러스 속(아르티큘라바이러스목; 오르토믹소바이러스과), 맘만타바이러스 아과, 바람직하게는 로안 바이러스, 모바트 바이러스, 오르토한타바이러스 또는 토팀바이러스 속(분야바이러스목; 한타바이러스과), 또는 뉴모바이러스과 속(모노네가바이러스목, 파라마이옥소바이러스과)의 바이러스일 수 있다.
따라서, 본원에 개시된 바와 같은 바이러스는 H1N1-, H1N2-, H2N2-, H3N2-, H5N1-, H6N1-, H7N2-, H7N3-, H7N7-, H7N9, H9N2-, H10N7-, H10N8- 또는 H5N1-서브타입(아르티큘라바이러스목; 오르토믹소바이러스과, 알파인플루엔자바이러스 속), 퓨말라 바이러스, 신 놈브레 바이러스, 서울 바이러스, 한탄 바이러스, 도브라바-벨그라드 바이러스, 사레마 바이러스, 포 코너즈 바이러스 또는 안데스 바이러스(분야바이러스목; 한타바이러스과, 맘만타바이러스과 아과, 오르토한타바이러스 속) 또는 메타뉴모바이러스, 오르토뉴모바이러스, 예컨대, 인간 오르토뉴모바이러스, 예컨대, 인간 호흡기 세포융합 바이러스 A, 인간 호흡기 세포융합 바이러스 B 또는 미분류 인간 호흡기 세포융합 바이러스, 개 뉴모바이러스, 고양이 뉴모바이러스, 양 호흡기 세포융합 바이러스, 양 호흡기 세포융합 바이러스(균주 WSU 83-1578), 뉴모바이러스, 호흡기 세포융합 바이러스, 돼지 뉴모바이러스 또는 뉴모바이러스 종(모노네가바이러스목, 파라마이옥소바이러스과, 뉴모바이러스과 속)일 수 있다.
ROCK 억제제는 ROCK 억제제의 투여 전 바이러스 부하와 비교하여 바이러스 부하를 감소시키는 것으로 구상된다. 예를 들어, 바이러스 부하는 플라크 형성 단위(pfu)/ml로 측정함으로써 결정될 수 있다.
이러한 맥락에서, "바이러스 부하를 감소시키는"은 mL 당 바이러스 입자 또는 감염성 입자가 ROCK 억제제의 투여 전에 존재하는 감염성 입자와 비교하여 적어도 10%, 20%, 30%, 40%, 50%, 60%, 70%, 80%, 90%, 95%, 99% 또는 100% 감소되는 것을 의미할 수 있다.
추가로, ROCK 억제제는 ROCK 억제제의 투여 전에 존재하는 폐의 체액 중량과 비교하여 폐의 체액 중량을 감소시킬 수 있는 것으로 고려된다. 체액 중량을 측정할 수 있는 방법은 실시예에 기재되어 있다.
또한, ROCK 억제제는 ROCK 억제제의 투여 전에 대식세포의 폐로의 침윤과 비교하여 폐로의 대식세포의 침윤을 감소시키는 것으로 고려된다. 대식세포의 침윤을 측정할 수 있는 방법은 실시예에 기재되어 있다.
본 발명은 또한 폐 상피 세포에서 정단 NKA 국소화를 방지하거나 감소시키는 데 효과적인 억제제의 결정을 위한 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스, 바람직하게는 인플루엔자 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템의 용도에 관한 것이다. 바람직하게는, 억제제는 본원에 기재된 바와 같은 ROCK 억제제이다.
시험 시스템에서 시험된 억제제는 시험관내 시험 시스템과 억제제를 접촉시킬 때 접촉 전 시험관내 시험 시스템에 존재하는 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킨다.
시험관내 시험 시스템은 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스, 바람직하게는 인플루엔자 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 임의의 적합한 시험관내 시험 시스템일 수 있다. 인플루엔자 바이러스는 균주 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)일 수 있다. 배양된 세포는, 예를 들어, 실시예 1에 기재된 바와 같이 배양된 Calu-3 세포일 수 있다.
배양된 폐 상피 세포는 인간 폐 상피 세포일 수 있다. 폐 상피 세포는 배지에 약 1×105, 2×105, 3×105, 4×105, 5×105, 6×105, 7×105, 8×105, 9×105, 10×105, 11×105의 밀도로 시딩될 수 있다.
본원에서 사용되는 용어 "접촉하는"은 바이러스-감염 배양된 폐 상피 세포를 공간적으로 관심 억제제에 매우 근접하게 하는 것을 지칭한다. 이는, 예를 들어, 관심 억제제가 배양된 세포를 주사기를 통해 위치시킨 배지에 적용됨을 의미할 수 있다. 본원에 기재된 바와 같이, 배양된 폐 상피 세포를 바이러스와 접촉시키는 단계는 억제제가 배양된 폐 상피 세포에 첨가되기 전에 수행된다.
따라서, 인플루엔자 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템의 사용은 시험 시스템을 시험하고자 하는 억제제(관심 억제제)와 접촉시키는 것을 포함할 수 있다.
억제제와 시험 시스템의 접촉이 억제제와 접촉하기 전에 존재하는 NKA의 정단 국소화와 비교하여 폐 상피 세포에서 NKA의 정단 국소화의 감소를 야기한다면, 억제제는 폐 상피 세포에서 NKA의 정단 국소화를 감소시키는 데 효율적이다.
ROCK 억제제는 시험관내 시험 시스템과 접촉할 때, 접촉 전 시험관내 시험 시스템과 비교하여, 폐 상피 세포에서 정단 NKA 국소화를 감소시키는 것으로 구상되며, 여기서 시험 시스템은 본원에 기재된 바와 같은 바이러스에 감염되는 배양된 폐 상피 세포를 포함한다.
본 발명은 또한
i)
폐 상피 세포에서 정단 NKA 국소화를 방지하거나,
ii)
ROCK 억제제의 투여 전 존재하는 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킴으로써,
폐부종의 예방 및/또는 치료 방법에 사용하기 위한 ROCK 억제제를 포함하는 조성물로서,
폐부종은 바이러스 감염과 관련되고, 바이러스는 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목인, 조성물에 관한 것이다.
ROCK 억제제를 포함하는 조성물은 약학적 조성물일 수 있다. 바람직하게는, 이러한 조성물은 담체, 바람직하게는 약학적으로 허용되는 담체를 추가로 포함한다. 조성물은 경구 투여 가능한 현탁액 또는 정제; 비강 스프레이, 멸균 주사용 제제(정맥내, 흉막내, 근육내), 예를 들어, 멸균 주사 가능한 수성 또는 유성 현탁액 또는 좌제의 형태일 수 있다. 현탁액으로서 경구 투여될 때, 이러한 조성물은 약학적 제형의 분야에서 이용 가능한 기술에 따라 제조되며, 당 분야에 공지된, 벌크를 제공하기 위한 미세결정질 셀룰로스, 현탁제로서 알긴산 또는 소듐 알기네이트, 점도 향상제로서 메틸셀룰로스, 및 감미제/향미제를 함유할 수 있다. 속방형 정제로서, 이러한 조성물은 미세결정질 셀룰로스, 디칼슘 포스페이트, 전분, 마그네슘 스테아레이트 및 락토스 및/또는 당 분야에 공지된 다른 부형제, 결합제, 증량제, 붕해제, 희석제, 및 윤활제를 함유할 수 있다. 주사용 용액 또는 현탁액은 적합한 비독성, 비경구적으로 허용되는 희석제 또는 용매, 예컨대, 만니톨, 1,3-부탄디올, 물, 링거액 또는 등장성 염화나트륨 용액, 또는 적합한 분산 또는 습윤제 및 현탁제, 예컨대, 합성 모노- 또는 디글리세리드를 포함하는 멸균, 순수, 고정 오일, 및 올레산을 포함하는 지방산을 사용하여 공지된 기술에 따라 제형화될 수 있다. 억제제 또는 억제제들은 바람직하게는 치료적 유효량으로 투여된다.
본 발명은 또한 폐 상피 세포에서 RNA 바이러스 관련 정단 NKA 국소화를 방지함으로써 폐부종을 갖거나 이의 위험이 있는 대상체를 치료하는 방법에 관한 것이다.
본원에서 사용되는 "대상체"는 임의의 적합한 대상체일 수 있다. 바람직하게는, 본원에서 사용되는 용어 "대상체"는 포유동물을 지칭한다. 대상체는 개, 고양이, 말, 양, 염소, 소 또는 인간 대상체, 바람직하게는 인간 대상체일 수 있다. 대상체는 본원에 기재된 바와 같은 폐부종을 갖는 대상체일 수 있다. 대상체는 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스에 감염된 대상체일 수 있다. 바람직하게는, 대상체는 오르토믹소바이러스속, 뉴모바이러스속 및/또는 한타바이러스과의 하나 이상의 바이러스에 감염된 대상체이다. 예를 들어, 이는 인플루엔자 A 바이러스와 같은 인플루엔자 바이러스에 감염된 대상체일 수 있다.
본원에서 사용되는 "감염 위험이 있는 대상체"는 본원에 기재된 바와 같이 폐부종이 발병할 위험이 있는 대상체일 수 있다. 대상체는 아르티큘라바이러스 목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스에 감염 위험이 있는 대상체일 수 있다. 바람직하게는, 대상체는 오르토믹소바이러스과, 뉴모바이러스과 및/또는 한타바이러스과 중 하나 이상의 바이러스에 감염 위험이 있는 대상체이다. 예를 들어, 이는 인플루엔자 A 바이러스와 같은 인플루엔자 바이러스에 감염 위험이 있는 대상체일 수 있다.
추가로, 대상체는 오르토믹소바이러스과(아르티큘라바이러스목), 아레나바이러스과, 한타바이러스과, 미포바이러스과, 나이로바이러스과, 페리분야바이러스과, 페뉴바이러스과(분야바이러스목), 보르나바이러스과, 파라바이러스과 및/또는 순바이러스과(모노네가바이러스목)의 바이러스에 감염된 대상체 또는 이로의 감염 위험이 있는 대상체인 것으로 구상된다.
또한, 대상체는 알파인플루엔자바이러스, 베타인플루엔자바이러스, 델타인플루엔자바이러스, 감마인플루엔자바이러스, 바람직하게는 알파인플루엔자바이러스 속(아르티큘라바이러스목; 오르토믹소바이러스과), 맘만타바이러스과 아과, 바람직하게는 로안 바이러스, 모바트 바이러스, 오르토한타바이러스 또는 토팀바이러스 속(분야바이러스목; 한타바이러스과), 또는 뉴모바이러스 속(모노네가바이러스목, 파라마이옥소바이러스과)의 바이러스에 감염된 대상체 또는 이로의 감염 위험이 있는 대상체인 것으로 고려된다.
추가로, 대상체는 H1N1-, H1N2-, H2N2-, H3N2-, H5N1-, H6N1-, H7N2-, H7N3-, H7N7-, H7N9, H9N2-, H10N7-, H10N8- 또는 H5N1-서브타입(아르티큘라바이러스 목; 오르토믹소바이러스과, 알파인플루엔자바이러스 속), 퓨말라 바이러스, 신 놈브레 바이러스, 서울 바이러스, 한탄 바이러스, 도브라바-벨그라드 바이러스, 사레마 바이러스, 포 코너즈 바이러스 또는 안데스 바이러스(분야바이러스목; 한타바이러스과, 맘만타바이러스과 아과, 오르토한타바이러스 속) 또는 메타뉴모바이러스, 오르토뉴모바이러스, 예컨대, 인간 오르토뉴모바이러스, 예컨대, 인간 호흡기 세포융합 바이러스 A, 인간 호흡기 세포융합 바이러스 B 또는 미분류 인간 호흡기 세포융합 바이러스, 개 뉴모바이러스, 고양이 뉴모바이러스, 양 호흡기 세포융합 바이러스, 양 호흡기 세포융합 바이러스(균주 WSU 83-1578), 뉴모바이러스, 호흡기 세포융합 바이러스, 돼지 뉴모바이러스 또는 뉴모바이러스 종(모노네가바이러스목, 파라마이옥소바이러스과, 뉴모바이러스속)에 감염된 대상체 또는 이로의 감염 위험이 있는 대상체인 것으로 구상된다.
또한, 대상체는 본원에 개시된 바와 같은 2종 이상의 바이러스에 감염된 대상체인 것으로 고려된다.
본 발명은 또한 폐부종의 예방 및/또는 치료에 효과적인 분자를 검출하는 방법으로서, 관심 화합물과 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스, 바람직하게는 인플루엔자 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템을 접촉시키는 것을 포함하고, 관심 화합물은 접촉 전의 시험관내 시험 시스템과 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시키는, 방법에 관한 것이다.
관심 화합물은 본원에 기재된 바와 같은 ROCK 억제제일 수 있다. 따라서, 상기 방법은 (i) 본원에 기재된 바와 같은 시험관내 시험 시스템에서 아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스, 바람직하게는 인플루엔자 바이러스에 감염된 폐 상피 세포에서 NKA 세포 국소화를 검출하는 단계 (ii) 시험관내 시험 시스템을 관심 화합물과 접촉시키는 단계, (iii) 시험관내 시험 시스템을 단계 (iii)의 관심 화합물과 접촉시킨 후 폐 상피 세포에서 NKA 세포 국소화를 검출하는 단계 및 (iv) 접촉 후 시험관내 시험 시스템에서 NKA의 국소화(즉, 이동)를 분석하는 단계로서, 접촉 전 정단 NKA 국소화와 비교하여 정단 NKA 국소화의 감소는 억제제가 폐부종의 예방 및/또는 치료에 효과적임을 지시하는 단계를 포함할 수 있다.
본 발명은 또한
i)
ROCK 억제제;
ii)
폐 상피 세포;
iii)
아르티큘라바이러스목, 모노네가바이러스목 및/또는 분야바이러스목의 바이러스; 및
iv)
NKA의 검출 및 세포 국소화를 위한 수단
을 포함하는 시험 시스템에 관한 것이다.
본원에서 사용되는 "NKA의 검출 및 세포 국소화를 위한 수단"은 임의의 적합한 수단일 수 있다. 이러한 수단은 당업자에게 공지되어 있다. 예를 들어, 당업자는 실시예에 기재된 바와 같은 수단을 사용할 수 있다.
하기 서열은 본 개시에서 언급되었다.
실시예
하기 실시예는 본 발명을 예시한다. 이러한 실시예는 본 발명의 범위를 제한하는 것으로 해석되어서는 안된다. 실시예는 예시의 목적으로 포함되며, 본 발명은 청구범위에 의해서만 제한된다.
실시예 1 - 세포 배양물 배양
모든 세포주를 5% CO2의 95% 가습 분위기 하에 37℃에서 75 cm2 또는 165 cm2 조직 배양 플라스크에서 배양하였다. 세포 단층이 90% 전면생장에 도달했을 때, 세포(최대 50%로 100% 전면생장에 도달하지 않은 Calu3 세포 제외)를 PBS -/-로 1회 세척하고 트립신-EDTA로 분리하였다. 세포를 적합한 배지(상기 표 참조)에 재현탁시키고, 각 실험 24 시간 전에 6-웰, 12-웰, 24-웰 조직 플레이트에서 배양 웰 내에 배치된 멸균 유리-커버 슬립 상에 또는 15 cm 조직 디쉬에 시딩하였다.
실시예 2 - Calu3 세포의 분극화
고도로 분극화된 Calu3 세포를 수득하기 위해, 50% 전면생장까지 생장된 175 cm2 조직 배양 플라스크에서 세포 단층을 PBS -/-로 1회 세척하고 트립신-EDTA로 처리하였다. 세포를 10 ml 배양 배지에 재현탁시키고, 300 xg, 24℃에서 15 min 동안 원심분리하였다. 상청액을 버리고 세포를 5 ml의 배양 배지에 재현탁시켰다. 30 μl의 세포 현탁액을 30 μl의 0.4% 트립판 블루 염료와 혼합하고, 세포 농도를 제조사의 지시에 따라 Neubauer 챔버에서 계산하였다. 세포를 Calu3 배양 배지에서 2×106개 생존 세포/ml의 농도로 희석하고, 0.5×106/0.25×106개 생존 세포를 함유하는 250 μl/125 μl의 세포 현탁액을 12/24 웰 플레이트에서 각각의 Transwells® 인서트의 정단 구획에 넣었다. 1/0.3 ml의 Calu-3 배양 배지를 기포의 혼입을 피하면서 기저측 구획에 첨가하고, 세포를 37℃, 5% CO2에서 배양하였다. 액체-액체 계면(LLI) 조건 하에 생장한 세포의 경우, 2일 마다 둘 모두의 구획에서 배지를 교체하였다. 공기-액체 계면(ALI)의 경우, 배양 배지를 2일 마다 정단 구획으로부터 흡인한 반면, 배지를 기저측 구획에서 2일 마다 교체하였다.
실시예 3 - 세포 생존율 검정
적용된 억제제의 세포독성을 확인하기 위해 상업적으로 이용 가능한 시약 PrestoBlue™를 사용하였다. 시약 - 레자주린(7-하이드록시-10-옥시도페녹사진-10-윰-3-온)-기반 화합물은 색 변화와 함께 생존 세포의 미토콘드리아 효소에 의해 환원된 형태로 전환되고, 분광광도계 또는 형광 접근법을 이용하여 정량화될 수 있다. 생존율 검정을 제조사의 프로토콜에 따라 수행하였다. Calu3 세포를 90 μl의 배양 배지에서 1 × 104/웰의 농도로 96-웰 플레이트에 시딩하고, 24 h 후에 상이한 농도의 억제제를 함유하는 배지로 처리하고, 24 h 후에 10 μl의 10-배 즉시 사용 가능한 PrestoBlue™ 시약을 각 웰에 첨가하였다. 이어서, 플레이트를 암실 하에 37℃에서 30 min 인큐베이션한 후, 흡광도를 Tecan Spark® 10 M 다중모드 마이크로플레이트 리더에 의해 570 nm 파장에서 측정하여 레자주린 전환의 양을 결정하였다.
실시예 4 - 벡터 물 수송
Calu3 세포 단층의 생리학적 상태의 특성으로서 벡터 물 수송(VWT)을 액체-액체 계면 조건 하에 14일 동안 Transwells® 인서트 상에서 생장한 분극화된 Calu3 세포의 정단 및 기저 세포 배양 배지에서 FITC-덱스트란 농도의 변화에 의해 측정하였다. 이를 위해, 세포를 모의 감염시키거나 감염 다중도(MOI): 2에서 37℃에서 1 시간 동안 PR8에 감염시켰다. 접종물을 제거하고 세포에 DMSO 또는 정확히 동일한 양의 DMSO(용매) 중 1 mg/ml의 70 kDa FITC-덱스트란 및 5 μM의 Rho-키나제 억제제 XIII를 함유하는 감염 배지 #2를 공급하였다. 세포를 37℃에서 8 및 24 h 동안 인큐베이션하였다. 정단 및 기저 쪽으로부터 30 μl의 세포 배양 배지를 수집하고, PBS -/-로 1:1 희석하고, 96 웰 평저형 검정색 플레이트에 놓았다. 샘플의 형광 강도를 Tecan Spark® 10M 다중모드 마이크로플레이트 리더에 의해 여기 파장 480 nm 및 방출 파장 535 nm에서 측정하였다. VWT는 하기 식을 이용하여 계산하였다:
여기서,
C0 - 출발점에서 배양 배지의 형광 값;
Ca - Transwells® 인서트의 정단 쪽에서 배양 배지의 형광 값;
Cb - Transwells® 인서트의 기저 쪽에서 배양 배지의 형광 값.
IAV 감염 동안 NKA 오분포가 IAV 병원성의 일반적 특징인지 여부를 결정하기 위해, 상이한 IAV 서브타입을 정단 NKA 발현을 유도하는 이들의 능력에 대해 스크리닝하였다. 먼저, Calu3 세포에서 인플루엔자 바이러스 A/Victoria/3/75 (H3N2), A/태국/1 (KAN-1)/2004 (H5N1) 또는 A/안휘/1/2013 (H7N9)의 생장 동역학을 비교하였다. 분석된 IAV 서브타입 및 이전에 시험된 PR8 바이러스의 복제 효율에는 유의한 차이가 검출되지 않았다. H5N1- 및 H7N9 서브타입의 고병원성 바이러스는 48 h p.i.에 각각 6.1 log10 FFU/ml 및 7.5 log10 FFU/ml와 동일한 최대 바이러스 역가를 나타낸 반면, 덜 병원성인 H3N2 균주는 24 h p.i.에 6.9 log10 FFU/ml의 최대 역가에 도달하였다(도 11a). 모든 시험된 바이러스는 OCWB 분석에 의해 입증된 바와 같이 바이러스 감염의 후기 단계에서 정단 NKA 불국소화를 유도할 수 있었다(도 11b). 상이한 IAV 균주 간에 정단 NKA 출현을 야기하는 능력에는 유의한 차이가 없었는데, 이는 이것이 분극화된 상피 세포의 IAV 감염에 의해 초래된 일반적인 효과임을 지시한다.
실시예 5 - 바이러스 전파
인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)를 165 cm2 배양 플라스크에서 MDCK II 세포에서 전파시켰다. 이를 위해 24-hr된 85% 전면생장된 단층 세포를 PBS -/-로 1회 세척하고, 0.01과 동일한 MOI에 상응하는 5 ml의 PBS +/+/BA/PS 함유 바이러스 희석액을 첨가한 후, 45 min 동안 실온에서 인큐베이션하였다. 후속하여, 접종물을 제거하고, 세포를 PBS -/-로 세척하고, 1 mg의 TPCK-처리된 트립신 ml-1을 함유하는 감염 배지 #1에서 37℃에서 2일 동안 인큐베이션하였다. 상청액을 수집하고 바이러스 역가를 병소-형성 검정에 의해 결정하였다.
병소 형성 검정
병소 형성 검정을 위해, MDCK II 세포를 3×106개 세포/플레이트의 농도로 96-웰 플레이트에 시딩하였다. 다음날, PBS +/+/BA/PS에서 이중으로 10배 희석액(10-1 내지 10-8)을 U-자형 96-웰의 각 바이러스 샘플로부터 제조하였다. 중요하게는, 희석액의 제조 동안 각 희석 단계 후에 피펫 팁을 변경하였다. 96-웰 플레이트(들)에서 MDCK II 세포를 PBS+/+로 1회 세척하였다. 이어서, U-자형 플레이트의 각 샘플에 대한 50 μl의 해당 희석액을 96-웰 플레이트의 해당 웰에 있는 MDCK II 세포로 옮긴 후, 이를 45 min 동안 인큐베이션하였다. 인큐베이션 후, 피펫 팁을 변경하지 않고 10-8 희석 열로 시작하여 접종물을 제거하고, 1 mg TPCK-처리 트립신 ml-1을 함유하는 100 μl의 아비셀(Avicel) 배지를 각 웰에 첨가하였다. 세포를 37℃, 5% CO2에서 30 시간 동안 인큐베이션한 후 면역세포화학 분석을 통해 바이러스-감염된 세포를 검출하였다. 이를 위해, 세포를 200 μl의 PBS +/+로 2회 세척하고, 고정시키고, 실온에서 30 분 동안 1%(v/v) Triton-X-100을 함유하는 4%(w/v) 파라포름알데하이드(PFA)에 투과시켰다. 다음으로, 세포를 400 μl의 세척 완충제(0.05% (v/v) Tween® 20이 있는 PBS +/+)로 3회 세척하고, 50 μl의 일차 항-NP 항체 용액(PBS +/+ 중 3% (w/v)의 BSA)으로 2 시간 동안 실온에서 덮었다. 이후, 세포를 세척 완충제로 3회 세척한 후, 50 μl의 이차 호스-래디쉬 퍼옥시다제(HRP) 표지된 항-마우스 항체와 함께 인큐베이션하였다. 1 시간 후, 세포를 다시 400 μl의 세척 완충제로 세척하고, 40 μl의 3-아미노-9-에틸카르바졸(AEC)-염색 완충제(아세테이트 완충제(50 mM 암모늄 아세테이트, 8.8 mM H2O2) 중 1x AEC 희석된 NN-디메틸포름아미드)를 각각의 웰에 첨가하였다. 병소가 검출될 수 있을 때까지 37℃에서 30 min 동안 인큐베이션한 후, 염색 완충제를 제거하고 세포를 dH2O로 2회 세척하였다. 공기-건조된 플레이트를 1200 dpi에서 Epson Perfection V500 Photo scan(Epson)을 사용하여 스캔하고, 웰 당 병소의 총 수를 결정하였다. 아비셀-배지는 주변 배지에서 바이러스 입자의 확산을 방지하는 높은 점도를 갖기 때문에, 바이러스는 한 세포에서 다른 형성 병소로만 확산될 수 있다.
1 ml 당 바이러스 역가는 하기 식에 의해 결정하였다:
여기서, fffu는 병소 형성 단위이다.
바이러스 적정을 위한 폐 균질물의 제조
병소 검정에 의한 감염된 폐 상피 세포에서 바이러스 역가의 분석을 위해, 마우스를 방혈에 의해 희생시켰다. 폐 순환을 우심실을 통해 멸균 PBS로 플러싱하였다. 플러싱된 블랜칭된 폐를 제거하고 차가운 PBS -/-로 세척하였다. 돌출부를 가위로 절단하고, 나머지 조직을 1 ml의 PBS -/-에서 단일 세포 현탁액으로 피펫팅함으로써 해리시켰다. 세포를 4℃에서 10 min 동안 400 xg에서 원심분리에 의해 펠렛화하고, 상청액을 앞서 기재된 바와 같이 병소 검정에 적용하였다.
실시예 6 - 면역형광 검정을 위한 세포의 고정
면역형광 검정을 위해 세포를 PBS +/+로 1회 세척하고 지시된 시점에 추가 투과의 존재 또는 부재 하에 고정시켰다. 사용된 일차 항체에 따라 세포를 고정시키고, 유기 용매로 투과시키거나 가교제 파라포름알데히드로 고정시켰다. 유기 용매로서 미리 냉각된(-20℃) 1:1(v/v) 아세톤:메탄올 용액(NKA α1 염색용) 또는 미리 냉각된(-20℃) 100% 메탄올(튜불린 염색용)을 -20℃에서 3 min 동안 사용한 후, 세척 완충제(0.05% (v/v) Tween® 20이 있는 PBS +/+)로 3회 세척하고, 차단 완충제(1x PBS +/+ 중 소 혈청 알부민(BSA) 3%(w /v))로 RT에서 1 시간 동안 또는 4℃에서 밤새 차단하였다. 가교 시약으로서 4%(w/v) PFA 용액을 사용하여 RT에서 10 min 동안 세포를 고정시킨 다음, 30 mM의 글리신을 함유하는 PBS +/+(G-PBS)로 3회 세척한 다음, 0.25% (v/w)의 Triton X-100으로 7 min 동안 투과시켰다. 이후, 세포를 G-PBS로 3회 세척하고, RT에서 30 min 동안 차단 용액(G-PBS 중 3% (w/v) BSA, G-PBS/BSA)으로 덮었다. 이후, 고정된 세포를 G-PBS 중 0.25% (v/w)의 Triton X-100으로 15 분 동안 처리하고, G-PBS로 3회 세척한 후, G-PBS/BSA에서 30 min 동안 차단하였다.
실시예 7 - 감염 및 비감염 Calu3 세포에서 정단 NKAα1 국소화의 항체 염색 및 공초점 레이저-스캐닝 현미경(도 1)
공기/액체-간기에서 Transwell® 인서트 상에서 생장한 Calu3 세포의 고 분극화된 단층을 감염되지 않은 채로 두거나 MOI = 5에서 인플루엔자 바이러스 A/푸에르토리코/8/34(H1N1)에 감염시켰다. 20 h p.i.에 감염된 세포를 처리하지 않은 채로 두거나 ROCK 억제제(XIII)로 처리하였다. 면역형광 검정을 위해 세포를 PBS +/+로 1회 세척하였다. 세포를 고정시키고, -20℃에서 3 min 동안 사전-냉각된(-20℃) 1:1(v/v) 아세톤:메탄올 용액(NKAα1 염색용)으로 투과시키고, 이어서 세척 완충제(1x PBS +/+ 중 소 혈청 알부민 0.3% (w/v))으로 3회 세척하고, 차단 완충제(1x PBS +/+ 중 소 혈청 알부민 3% (w/v))로 RT에서 1 시간 동안 또는 4℃에서 밤새 차단하였다. 이후 항체 염색을 위해 세포를 항체 희석 용액(1x PBS +/+ 중 소 혈청 알부민 2% (w/v)) 중 특정 일차 항체(토끼 항-NP, Thermo-Fisher(PA5-32242): 1:2000; 마우스 항-α1 NKA, Millipore/Sigma Aldrich (# 05-369): 1:1000)와 인큐베이션하였다. 항체 희석제를 RT에서 2 h 동안 고정되고 투과된 세포에 첨가한 다음, PBS +/+로 2회 세척하였다. 이후, 세포를 항체 희석 용액(1:1000)에 희석된 이차 항체(닭 항-토끼 Alexa Fluor 488 및 닭 항-마우스 Alexa Fluor 647)와 함께 1 h 동안 인큐베이션하였다. 이후, 세포를 PBS -/-로 3회, ddH2O로 1회 세척하고, Transwells®로부터의 커버 슬립 또는 폴리에스테르 막을 DAPI(농도는 제조사에 의해 제공되지 않음)를 갖는 ProLong™ Gold 변색 방지제와 함께 유리 슬라이드 상에 밤새 장착하였다. 간접 면역형광 분석 및 Imaris® 소프트웨어를 사용한 후속 3D-모델링에 의해 NKAα1 국소화를 평가하였다. HCX PL Apo 63x/1.30 GLYC 대물렌즈 및 핀홀 - 1 에어리 유닛(AU)을 구비한 Leica TCS-SP5 공초점 레이저-스캐닝 현미경을 사용하여 신호를 가시화하였다. Z-스택을 0.25 μm 스텝 크기를 사용하여 획득하고, 결과를 Imaris 소프트웨어(Bitplane)에 의해 분석하였다(도 1 및 2).
실시예 8 - 단백질 세포 표면 발현의 분석
온-셀-웨스턴 블롯 검정
온-셀 웨스턴 블롯 검정을 위해, 3 세트의 Calu3 세포를 6×104개 세포/웰의 농도로 광학적 투명 평저형 플레이트가 있는 96-웰에 시딩하였다. 24 시간 후, 세포 단층이 95% 전면생장일 때, 세포를 2의 MOI에서 인플루엔자 바이러스 A/푸에르토리코78/34(H1N1)에 감염시켰다. 37℃에서 45 min 인큐베이션 후, 접종물을 감염 배지(1% 소듐 피루베이트(100x), 1% 비필수 아미노산(100x), 0.5% BSA(30%)를 함유하는 MEM) (+/-) 억제제(파수딜 HCl, Selleckchem: 10 μM; Rho 키나제 억제제 RKI-1447(XIII), Millipore: 5 μM) 또는 대조로서 억제제의 용매로 교체하였다. 24 시간 후 억제제 또는 용매를 함유하는 배지를 제거하고, PBS +/+에 희석된 HA, M2 또는 Na+,K+-ATPase β1 서브유닛의 세포외 에피토프를 인식하는 일차 항체(염소 다클론성 항-인플루엔자 A 바이러스, Abcam(# ab20841): 1:2000; 마우스 단클론성 항-인플루엔자 A 바이러스 M2, ThermoScintific/Invitrogen/Gibco (# MA1-082): 1:1000; 마우스 단클론성 항-β1 NKA, ThermoScintific/Invitrogen/Gibco (# MA3-930): 1:1000)을 Calu3 세포의 한 세트에 첨가하고, 플레이트를 5% CO2에서 37℃에서 1.5 h 동안 추가로 인큐베이션하였다. 이후, 세포를 PBS+/+로 3회 세척하고, RT에서 20 min 동안 4% PFA로 고정시킨 다음, PBS +/+로 각각 5 min 동안 3회 세척하였다. 이후, 세포를 RT에서 45 min 동안 차단 완충제로 처리한 다음, RT에서 1 h 동안 5 μM의 DRAQ5™(원적색 DNA 염색제)을 함유하는 차단 완충제로 희석된 이차 IRDye 800-접합된 항-마우스, 또는 염소 항체(LI-COR, 일차 항체의 숙주에 따라)와 암실 하에 인큐베이션하였다. 이어서, 세포를 TBS-T로 3회 세척하고 LI-Cor Odyssay 적외선 이미저(100 μm 해상도, 0.5 mm 포커스 오프셋)에서 스캔하였다. Image Studio(LI-COR), 엑셀(Microsoft) 및 GraphPad Prism 5(Graphpad Software, Inc.) 소프트웨어를 사용하여 데이터를 분석하였다(도 11).
실시예 9 - 동물 생체내 실험
모든 동물 실험은 "유럽 실험 동물 과학 협회의 연합(Federation of European Laboratory Animal Science Associations; FELASA)"의 최신 지침에 따라 수행되었으며, 부에노스 아이레스의 생물의학 조사(Investigacion en Biomedicina de Buenos Aires; IBioBA)로부터 심장 및 폐 연구소의 맥스-플랑크 실험실(Max-Planck Laboratory)의 지역 위원회의 승인을 받았다. 6주령된 BALB/c 마우스(그룹당 n = 5)를 30 μl의 부피로 PR8의 500 플라크-형성 단위(pfu)/마우스의 기관내 접종에 의해 감염시켰다. 파수딜 HCl을 멸균 PBS -/-에 희석하고, 다음 7일 동안 10 mg/kg(24 h p.i.)의 농도로 매일 복강내(IP) 적용하였다. 대조로서 멸균 PBS의 IP 주사를 적용하였다. 체중을 8일차 p.i.까지 매일 모니터링하였다. 치료 시작 후 7일(= 8일차 p.i.)에 마우스를 과량의 이소플루란으로 희생시켰다.
습윤-대-건조 폐 중량 비율
폐 습윤-대-건조(W/D) 중량 비율을 이용하여 IAV 감염 후 폐수 축적을 분석하였다. 동물을 희생시키고, 해부하고, 절제 직후 폐 '습윤' 중량을 측정하였다. 이후, 폐를 60℃의 오븐에서 5일 동안 건조시키고 건조 중량으로서 재-칭량하였다. W/D 중량 비율은 습윤 중량을 건조 중량으로 나누어 계산하였다.
조직학적 처리를 위한 폐의 준비
동물을 희생시키고, 폐를 PBS -/-로 우심실을 통해 관류시키고, 흉강으로부터 제거하고, 24 h 동안 4% PFA에 고정시킨 다음, 파라핀(Leica ASP200S)에 포매하였다. 파라핀 포매된 폐를 Microtome RM2125(Leica)를 사용하여 얇은 절편(3.5 μm)으로 절단하였다. 슬라이스를 하전된 슬라이드에 장착하고 37℃에서 밤새 건조시켰다. 다음날, 폐 절편을 하기 절차에 의해 헤마톡실린/에오신에 의해 염색하였다:
본원에 기재된 모든 현미경 분석은 EVOS FL Auto Cell Imaging System에 의해 수행되었다. 조직학적 절단에서 세포의 총량은 Univ.-Prof. Dr. Achim Gruber (
)의 협력 그룹에서 "Aperio v9 핵 계수 알고리즘" 소프트웨어(Leica Biosystems Imaging Inc., CA, USA)를 사용하여 Aperio CS2 스캐너(Leica Biosystems Imaging Inc., CA, USA)에 의해 정량화하였다.
통계
통계 분석은 GraphPad Prism 5 소프트웨어에 의해 수행되었다. 데이터는 평균 + 평균의 표준 오차(SEM) 또는 평균의 표준 편차(SD)로 제공된다(도면 범례에 표시됨). 2개의 그룹의 통계적 유의성은 양측 비쌍형 스튜던트 t 검정에 의해 시험되었다. 3개 이상의 그룹의 통계적 유의성은 일원 ANOVA 및 이어서 터키(Tukey)의 사후 검정에 의해 분석되었다. P 값은 0.05 미만(*p<0.05; **p<0.01; ***p<0.005)이면 유의한 것으로 간주되었다.
***
달리 언급되지 않는 한, 설명 및 청구범위를 포함하여 본 문서에서 사용되는 하기 용어는 하기에 주어진 정의를 갖는다.
당업자는 단지 일상적인 실험을 이용하여 본원에 기재된 본 발명의 특정 실시양태에 대한 많은 등가물을 인지하거나 확인할 수 있을 것이다. 이러한 등가물은 본 발명에 포함되는 것으로 의도된다.
본원에 사용되는 단수 형태인 부정관사 및 정관사는 달리 분명하게 표시되지 않으면 복수의 대상을 포함한다는 것이 주지되어야 한다. 따라서, 예를 들어, "시약"에 대한 언급은 하나 이상의 이러한 상이한 시약을 포함하고, "방법"에 대한 언급은 본원에 기재된 방법을 변형 또는 치환할 수 있는 당해 기술분야의 숙련가에게 공지된 등가의 단계 및 방법에 대한 언급을 포함한다.
달리 명시하지 않는 한, 일련의 요소에 선행하는 용어 "적어도"는 일련의 모든 요소를 지칭하는 것으로 이해되어야 한다. 당업자는 단지 일상적인 실험을 이용하여 본원에 기재된 본 발명의 특정 실시양태에 대한 많은 등가물을 인지하거나 확인할 수 있을 것이다. 이러한 등가물은 본 발명에 포함되는 것으로 의도된다.
본원에서 사용되는 용어 "및/또는"은 "및", "또는" 및 "상기 용어에 의해 연결된 요소들의 전부 또는 임의의 다른 조합"의 의미를 포함한다.
본원에서 사용되는 용어 "약" 또는 "대략"은 주어진 값 또는 범위의 20% 이내, 바람직하게는 10% 이내, 및 더욱 바람직하게는 5% 이내를 의미한다. 그러나, 이는 또한 구체적인 수를 포함하며, 예를 들어, 약 20은 20을 포함한다.
본 명세서 및 하기 특허청구범위 전반에 걸쳐, 문맥이 달리 요구하지 않는 한, 단어 "포함하다" 및 변형어, 예컨대, "포함한다" 및 "포함하는"은 명시된 정수 또는 단계 또는 정수 또는 단계의 그룹의 포함을 의미하고 임의의 다른 정수 또는 단계 또는 정수 또는 단계의 그룹의 배제를 의미하는 것은 아닌 것으로 이해될 것이다. 본원에서 사용되는 용어 "포함하는"은 용어 "함유하는" 또는 "비롯한"으로 치환될 수 있거나, 본원에서 사용되는 경우에 종종 용어 "갖는"으로 치환될 수 있다.
본원에서 사용되는 "이루어진"은 특허청구항 요소에 지정되지 않은 임의의 요소, 단계 또는 성분을 배제한다. 본원에서 사용되는 "본질적으로 이루어진"은 특허청구항의 기본 및 신규 특징에 실질적으로 영향을 미치지 않는 재료 또는 단계를 배제하지 않는다.
본원의 각각의 예에서, 임의의 용어 "포함하는", "본질적으로 이루어진" 및 "이루어진"은 다른 2개 용어의 어느 하나로 대체될 수 있다
본 발명은 본원에 기재된 특정 방법, 프로토콜, 재료, 시약, 및 물질 등에 제한되지 않으며, 이에 따라 달라질 수 있음이 이해되어야 한다. 본원에서 사용되는 용어는 단지 특정 구현예를 설명하기 위한 것이며, 본 발명의 범위를 제한하려는 것이 아니고, 이는 특허청구범위에 의해서만 한정된다.
본 명세서의 텍스트 전반에 걸쳐 인용된 모든 간행물(모든 특허, 특허 출원, 과학 간행물, 제조자 명세서, 설명서 등을 포함)은 그 전체가 본원에 참조로 포함된다. 본원의 어떠한 것도 본 발명이 선행 발명에 의해 이러한 개시보다 선행할 자격이 없음을 인정하는 것으로 해석되어서는 안된다. 참조로 포함된 물질이 본 명세서와 모순되거나 불일치하는 정도까지, 본 명세서는 이러한 모든 물질을 대체할 것이다.
참고문헌 목록
SEQUENCE LISTING
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Gly Ser Val Lys Glu Met Arg Glu Arg Tyr Ala Lys Ile Val Glu Ile
465 470 475 480
Pro Phe Asn Ser Thr Asn Lys Tyr Gln Leu Ser Ile His Lys Asn Pro
485 490 495
Asn Thr Ser Glu Pro Gln His Leu Leu Val Met Lys Gly Ala Pro Glu
500 505 510
Arg Ile Leu Asp Arg Cys Ser Ser Ile Leu Leu His Gly Lys Glu Gln
515 520 525
Pro Leu Asp Glu Glu Leu Lys Asp Ala Phe Gln Asn Ala Tyr Leu Glu
530 535 540
Leu Gly Gly Leu Gly Glu Arg Val Leu Gly Phe Cys His Leu Phe Leu
545 550 555 560
Pro Asp Glu Gln Phe Pro Glu Gly Phe Gln Phe Asp Thr Asp Asp Val
565 570 575
Asn Phe Pro Ile Asp Asn Leu Cys Phe Val Gly Leu Ile Ser Met Ile
580 585 590
Asp Pro Pro Arg Ala Ala Val Pro Asp Ala Val Gly Lys Cys Arg Ser
595 600 605
Ala Gly Ile Lys Val Ile Met Val Thr Gly Asp His Pro Ile Thr Ala
610 615 620
Lys Ala Ile Ala Lys Gly Val Gly Ile Ile Ser Glu Gly Asn Glu Thr
625 630 635 640
Val Glu Asp Ile Ala Ala Arg Leu Asn Ile Pro Val Ser Gln Val Asn
645 650 655
Pro Arg Asp Ala Lys Ala Cys Val Val His Gly Ser Asp Leu Lys Asp
660 665 670
Met Thr Ser Glu Gln Leu Asp Asp Ile Leu Lys Tyr His Thr Glu Ile
675 680 685
Val Phe Ala Arg Thr Ser Pro Gln Gln Lys Leu Ile Ile Val Glu Gly
690 695 700
Cys Gln Arg Gln Gly Ala Ile Val Ala Val Thr Gly Asp Gly Val Asn
705 710 715 720
Asp Ser Pro Ala Leu Lys Lys Ala Asp Ile Gly Val Ala Met Gly Ile
725 730 735
Ala Gly Ser Asp Val Ser Lys Gln Ala Ala Asp Met Ile Leu Leu Asp
740 745 750
Asp Asn Phe Ala Ser Ile Val Thr Gly Val Glu Glu Gly Arg Leu Ile
755 760 765
Phe Asp Asn Leu Lys Lys Ser Ile Ala Tyr Thr Leu Thr Ser Asn Ile
770 775 780
Pro Glu Ile Thr Pro Phe Leu Ile Phe Ile Ile Ala Asn Ile Pro Leu
785 790 795 800
Pro Leu Gly Thr Val Thr Ile Leu Cys Ile Asp Leu Gly Thr Asp Met
805 810 815
Val Pro Ala Ile Ser Leu Ala Tyr Glu Gln Ala Glu Ser Asp Ile Met
820 825 830
Lys Arg Gln Pro Arg Asn Pro Lys Thr Asp Lys Leu Val Asn Glu Arg
835 840 845
Leu Ile Ser Met Ala Tyr Gly Gln Ile Gly Met Ile Gln Ala Leu Gly
850 855 860
Gly Phe Phe Thr Tyr Phe Val Ile Leu Ala Glu Asn Gly Phe Leu Pro
865 870 875 880
Ile His Leu Leu Gly Leu Arg Val Asp Trp Asp Asp Arg Trp Ile Asn
885 890 895
Asp Val Glu Asp Ser Tyr Gly Gln Gln Trp Thr Tyr Glu Gln Arg Lys
900 905 910
Ile Val Glu Phe Thr Cys His Thr Ala Phe Phe Val Ser Ile Val Val
915 920 925
Val Gln Trp Ala Asp Leu Val Ile Cys Lys Thr Arg Arg Asn Ser Val
930 935 940
Phe Gln Gln Gly Met Lys Asn Lys Ile Leu Ile Phe Gly Leu Phe Glu
945 950 955 960
Glu Thr Ala Leu Ala Ala Phe Leu Ser Tyr Cys Pro Gly Met Gly Val
965 970 975
Ala Leu Arg Met Tyr Pro Leu Lys Pro Thr Trp Trp Phe Cys Ala Phe
980 985 990
Pro Tyr Ser Leu Leu Ile Phe Val Tyr Asp Glu Val Arg Lys Leu Ile
995 1000 1005
Ile Arg Arg Arg Pro Gly Gly Trp Val Glu Lys Glu Thr Tyr Tyr
1010 1015 1020
<210> 5
<211> 1020
<212> PRT
<213> Homo sapiens
<400> 5
Met Gly Arg Gly Ala Gly Arg Glu Tyr Ser Pro Ala Ala Thr Thr Ala
1 5 10 15
Glu Asn Gly Gly Gly Lys Lys Lys Gln Lys Glu Lys Glu Leu Asp Glu
20 25 30
Leu Lys Lys Glu Val Ala Met Asp Asp His Lys Leu Ser Leu Asp Glu
35 40 45
Leu Gly Arg Lys Tyr Gln Val Asp Leu Ser Lys Gly Leu Thr Asn Gln
50 55 60
Arg Ala Gln Asp Val Leu Ala Arg Asp Gly Pro Asn Ala Leu Thr Pro
65 70 75 80
Pro Pro Thr Thr Pro Glu Trp Val Lys Phe Cys Arg Gln Leu Phe Gly
85 90 95
Gly Phe Ser Ile Leu Leu Trp Ile Gly Ala Ile Leu Cys Phe Leu Ala
100 105 110
Tyr Gly Ile Gln Ala Ala Met Glu Asp Glu Pro Ser Asn Asp Asn Leu
115 120 125
Tyr Leu Gly Val Val Leu Ala Ala Val Val Ile Val Thr Gly Cys Phe
130 135 140
Ser Tyr Tyr Gln Glu Ala Lys Ser Ser Lys Ile Met Asp Ser Phe Lys
145 150 155 160
Asn Met Val Pro Gln Gln Ala Leu Val Ile Arg Glu Gly Glu Lys Met
165 170 175
Gln Ile Asn Ala Glu Glu Val Val Val Gly Asp Leu Val Glu Val Lys
180 185 190
Gly Gly Asp Arg Val Pro Ala Asp Leu Arg Ile Ile Ser Ser His Gly
195 200 205
Cys Lys Val Asp Asn Ser Ser Leu Thr Gly Glu Ser Glu Pro Gln Thr
210 215 220
Arg Ser Pro Glu Phe Thr His Glu Asn Pro Leu Glu Thr Arg Asn Ile
225 230 235 240
Cys Phe Phe Ser Thr Asn Cys Val Glu Gly Thr Ala Arg Gly Ile Val
245 250 255
Ile Ala Thr Gly Asp Arg Thr Val Met Gly Arg Ile Ala Thr Leu Ala
260 265 270
Ser Gly Leu Glu Val Gly Arg Thr Pro Ile Ala Met Glu Ile Glu His
275 280 285
Phe Ile Gln Leu Ile Thr Gly Val Ala Val Phe Leu Gly Val Ser Phe
290 295 300
Phe Val Leu Ser Leu Ile Leu Gly Tyr Ser Trp Leu Glu Ala Val Ile
305 310 315 320
Phe Leu Ile Gly Ile Ile Val Ala Asn Val Pro Glu Gly Leu Leu Ala
325 330 335
Thr Val Thr Val Cys Leu Thr Leu Thr Ala Lys Arg Met Ala Arg Lys
340 345 350
Asn Cys Leu Val Lys Asn Leu Glu Ala Val Glu Thr Leu Gly Ser Thr
355 360 365
Ser Thr Ile Cys Ser Asp Lys Thr Gly Thr Leu Thr Gln Asn Arg Met
370 375 380
Thr Val Ala His Met Trp Phe Asp Asn Gln Ile His Glu Ala Asp Thr
385 390 395 400
Thr Glu Asp Gln Ser Gly Ala Thr Phe Asp Lys Arg Ser Pro Thr Trp
405 410 415
Thr Ala Leu Ser Arg Ile Ala Gly Leu Cys Asn Arg Ala Val Phe Lys
420 425 430
Ala Gly Gln Glu Asn Ile Ser Val Ser Lys Arg Asp Thr Ala Gly Asp
435 440 445
Ala Ser Glu Ser Ala Leu Leu Lys Cys Ile Glu Leu Ser Cys Gly Ser
450 455 460
Val Arg Lys Met Arg Asp Arg Asn Pro Lys Val Ala Glu Ile Pro Phe
465 470 475 480
Asn Ser Thr Asn Lys Tyr Gln Leu Ser Ile His Glu Arg Glu Asp Ser
485 490 495
Pro Gln Ser His Val Leu Val Met Lys Gly Ala Pro Glu Arg Ile Leu
500 505 510
Asp Arg Cys Ser Thr Ile Leu Val Gln Gly Lys Glu Ile Pro Leu Asp
515 520 525
Lys Glu Met Gln Asp Ala Phe Gln Asn Ala Tyr Met Glu Leu Gly Gly
530 535 540
Leu Gly Glu Arg Val Leu Gly Phe Cys Gln Leu Asn Leu Pro Ser Gly
545 550 555 560
Lys Phe Pro Arg Gly Phe Lys Phe Asp Thr Asp Glu Leu Asn Phe Pro
565 570 575
Thr Glu Lys Leu Cys Phe Val Gly Leu Met Ser Met Ile Asp Pro Pro
580 585 590
Arg Ala Ala Val Pro Asp Ala Val Gly Lys Cys Arg Ser Ala Gly Ile
595 600 605
Lys Val Ile Met Val Thr Gly Asp His Pro Ile Thr Ala Lys Ala Ile
610 615 620
Ala Lys Gly Val Gly Ile Ile Ser Glu Gly Asn Glu Thr Val Glu Asp
625 630 635 640
Ile Ala Ala Arg Leu Asn Ile Pro Met Ser Gln Val Asn Pro Arg Glu
645 650 655
Ala Lys Ala Cys Val Val His Gly Ser Asp Leu Lys Asp Met Thr Ser
660 665 670
Glu Gln Leu Asp Glu Ile Leu Lys Asn His Thr Glu Ile Val Phe Ala
675 680 685
Arg Thr Ser Pro Gln Gln Lys Leu Ile Ile Val Glu Gly Cys Gln Arg
690 695 700
Gln Gly Ala Ile Val Ala Val Thr Gly Asp Gly Val Asn Asp Ser Pro
705 710 715 720
Ala Leu Lys Lys Ala Asp Ile Gly Ile Ala Met Gly Ile Ser Gly Ser
725 730 735
Asp Val Ser Lys Gln Ala Ala Asp Met Ile Leu Leu Asp Asp Asn Phe
740 745 750
Ala Ser Ile Val Thr Gly Val Glu Glu Gly Arg Leu Ile Phe Asp Asn
755 760 765
Leu Lys Lys Ser Ile Ala Tyr Thr Leu Thr Ser Asn Ile Pro Glu Ile
770 775 780
Thr Pro Phe Leu Leu Phe Ile Ile Ala Asn Ile Pro Leu Pro Leu Gly
785 790 795 800
Thr Val Thr Ile Leu Cys Ile Asp Leu Gly Thr Asp Met Val Pro Ala
805 810 815
Ile Ser Leu Ala Tyr Glu Ala Ala Glu Ser Asp Ile Met Lys Arg Gln
820 825 830
Pro Arg Asn Ser Gln Thr Asp Lys Leu Val Asn Glu Arg Leu Ile Ser
835 840 845
Met Ala Tyr Gly Gln Ile Gly Met Ile Gln Ala Leu Gly Gly Phe Phe
850 855 860
Thr Tyr Phe Val Ile Leu Ala Glu Asn Gly Phe Leu Pro Ser Arg Leu
865 870 875 880
Leu Gly Ile Arg Leu Asp Trp Asp Asp Arg Thr Met Asn Asp Leu Glu
885 890 895
Asp Ser Tyr Gly Gln Glu Trp Thr Tyr Glu Gln Arg Lys Val Val Glu
900 905 910
Phe Thr Cys His Thr Ala Phe Phe Ala Ser Ile Val Val Val Gln Trp
915 920 925
Ala Asp Leu Ile Ile Cys Lys Thr Arg Arg Asn Ser Val Phe Gln Gln
930 935 940
Gly Met Lys Asn Lys Ile Leu Ile Phe Gly Leu Leu Glu Glu Thr Ala
945 950 955 960
Leu Ala Ala Phe Leu Ser Tyr Cys Pro Gly Met Gly Val Ala Leu Arg
965 970 975
Met Tyr Pro Leu Lys Val Thr Trp Trp Phe Cys Ala Phe Pro Tyr Ser
980 985 990
Leu Leu Ile Phe Ile Tyr Asp Glu Val Arg Lys Leu Ile Leu Arg Arg
995 1000 1005
Tyr Pro Gly Gly Trp Val Glu Lys Glu Thr Tyr Tyr
1010 1015 1020
<210> 6
<211> 1013
<212> PRT
<213> Homo sapiens
<400> 6
Met Gly Asp Lys Lys Asp Asp Lys Asp Ser Pro Lys Lys Asn Lys Gly
1 5 10 15
Lys Glu Arg Arg Asp Leu Asp Asp Leu Lys Lys Glu Val Ala Met Thr
20 25 30
Glu His Lys Met Ser Val Glu Glu Val Cys Arg Lys Tyr Asn Thr Asp
35 40 45
Cys Val Gln Gly Leu Thr His Ser Lys Ala Gln Glu Ile Leu Ala Arg
50 55 60
Asp Gly Pro Asn Ala Leu Thr Pro Pro Pro Thr Thr Pro Glu Trp Val
65 70 75 80
Lys Phe Cys Arg Gln Leu Phe Gly Gly Phe Ser Ile Leu Leu Trp Ile
85 90 95
Gly Ala Ile Leu Cys Phe Leu Ala Tyr Gly Ile Gln Ala Gly Thr Glu
100 105 110
Asp Asp Pro Ser Gly Asp Asn Leu Tyr Leu Gly Ile Val Leu Ala Ala
115 120 125
Val Val Ile Ile Thr Gly Cys Phe Ser Tyr Tyr Gln Glu Ala Lys Ser
130 135 140
Ser Lys Ile Met Glu Ser Phe Lys Asn Met Val Pro Gln Gln Ala Leu
145 150 155 160
Val Ile Arg Glu Gly Glu Lys Met Gln Val Asn Ala Glu Glu Val Val
165 170 175
Val Gly Asp Leu Val Glu Ile Lys Gly Gly Asp Arg Val Pro Ala Asp
180 185 190
Leu Arg Ile Ile Ser Ala His Gly Cys Lys Val Asp Asn Ser Ser Leu
195 200 205
Thr Gly Glu Ser Glu Pro Gln Thr Arg Ser Pro Asp Cys Thr His Asp
210 215 220
Asn Pro Leu Glu Thr Arg Asn Ile Thr Phe Phe Ser Thr Asn Cys Val
225 230 235 240
Glu Gly Thr Ala Arg Gly Val Val Val Ala Thr Gly Asp Arg Thr Val
245 250 255
Met Gly Arg Ile Ala Thr Leu Ala Ser Gly Leu Glu Val Gly Lys Thr
260 265 270
Pro Ile Ala Ile Glu Ile Glu His Phe Ile Gln Leu Ile Thr Gly Val
275 280 285
Ala Val Phe Leu Gly Val Ser Phe Phe Ile Leu Ser Leu Ile Leu Gly
290 295 300
Tyr Thr Trp Leu Glu Ala Val Ile Phe Leu Ile Gly Ile Ile Val Ala
305 310 315 320
Asn Val Pro Glu Gly Leu Leu Ala Thr Val Thr Val Cys Leu Thr Leu
325 330 335
Thr Ala Lys Arg Met Ala Arg Lys Asn Cys Leu Val Lys Asn Leu Glu
340 345 350
Ala Val Glu Thr Leu Gly Ser Thr Ser Thr Ile Cys Ser Asp Lys Thr
355 360 365
Gly Thr Leu Thr Gln Asn Arg Met Thr Val Ala His Met Trp Phe Asp
370 375 380
Asn Gln Ile His Glu Ala Asp Thr Thr Glu Asp Gln Ser Gly Thr Ser
385 390 395 400
Phe Asp Lys Ser Ser His Thr Trp Val Ala Leu Ser His Ile Ala Gly
405 410 415
Leu Cys Asn Arg Ala Val Phe Lys Gly Gly Gln Asp Asn Ile Pro Val
420 425 430
Leu Lys Arg Asp Val Ala Gly Asp Ala Ser Glu Ser Ala Leu Leu Lys
435 440 445
Cys Ile Glu Leu Ser Ser Gly Ser Val Lys Leu Met Arg Glu Arg Asn
450 455 460
Lys Lys Val Ala Glu Ile Pro Phe Asn Ser Thr Asn Lys Tyr Gln Leu
465 470 475 480
Ser Ile His Glu Thr Glu Asp Pro Asn Asp Asn Arg Tyr Leu Leu Val
485 490 495
Met Lys Gly Ala Pro Glu Arg Ile Leu Asp Arg Cys Ser Thr Ile Leu
500 505 510
Leu Gln Gly Lys Glu Gln Pro Leu Asp Glu Glu Met Lys Glu Ala Phe
515 520 525
Gln Asn Ala Tyr Leu Glu Leu Gly Gly Leu Gly Glu Arg Val Leu Gly
530 535 540
Phe Cys His Tyr Tyr Leu Pro Glu Glu Gln Phe Pro Lys Gly Phe Ala
545 550 555 560
Phe Asp Cys Asp Asp Val Asn Phe Thr Thr Asp Asn Leu Cys Phe Val
565 570 575
Gly Leu Met Ser Met Ile Asp Pro Pro Arg Ala Ala Val Pro Asp Ala
580 585 590
Val Gly Lys Cys Arg Ser Ala Gly Ile Lys Val Ile Met Val Thr Gly
595 600 605
Asp His Pro Ile Thr Ala Lys Ala Ile Ala Lys Gly Val Gly Ile Ile
610 615 620
Ser Glu Gly Asn Glu Thr Val Glu Asp Ile Ala Ala Arg Leu Asn Ile
625 630 635 640
Pro Val Ser Gln Val Asn Pro Arg Asp Ala Lys Ala Cys Val Ile His
645 650 655
Gly Thr Asp Leu Lys Asp Phe Thr Ser Glu Gln Ile Asp Glu Ile Leu
660 665 670
Gln Asn His Thr Glu Ile Val Phe Ala Arg Thr Ser Pro Gln Gln Lys
675 680 685
Leu Ile Ile Val Glu Gly Cys Gln Arg Gln Gly Ala Ile Val Ala Val
690 695 700
Thr Gly Asp Gly Val Asn Asp Ser Pro Ala Leu Lys Lys Ala Asp Ile
705 710 715 720
Gly Val Ala Met Gly Ile Ala Gly Ser Asp Val Ser Lys Gln Ala Ala
725 730 735
Asp Met Ile Leu Leu Asp Asp Asn Phe Ala Ser Ile Val Thr Gly Val
740 745 750
Glu Glu Gly Arg Leu Ile Phe Asp Asn Leu Lys Lys Ser Ile Ala Tyr
755 760 765
Thr Leu Thr Ser Asn Ile Pro Glu Ile Thr Pro Phe Leu Leu Phe Ile
770 775 780
Met Ala Asn Ile Pro Leu Pro Leu Gly Thr Ile Thr Ile Leu Cys Ile
785 790 795 800
Asp Leu Gly Thr Asp Met Val Pro Ala Ile Ser Leu Ala Tyr Glu Ala
805 810 815
Ala Glu Ser Asp Ile Met Lys Arg Gln Pro Arg Asn Pro Arg Thr Asp
820 825 830
Lys Leu Val Asn Glu Arg Leu Ile Ser Met Ala Tyr Gly Gln Ile Gly
835 840 845
Met Ile Gln Ala Leu Gly Gly Phe Phe Ser Tyr Phe Val Ile Leu Ala
850 855 860
Glu Asn Gly Phe Leu Pro Gly Asn Leu Val Gly Ile Arg Leu Asn Trp
865 870 875 880
Asp Asp Arg Thr Val Asn Asp Leu Glu Asp Ser Tyr Gly Gln Gln Trp
885 890 895
Thr Tyr Glu Gln Arg Lys Val Val Glu Phe Thr Cys His Thr Ala Phe
900 905 910
Phe Val Ser Ile Val Val Val Gln Trp Ala Asp Leu Ile Ile Cys Lys
915 920 925
Thr Arg Arg Asn Ser Val Phe Gln Gln Gly Met Lys Asn Lys Ile Leu
930 935 940
Ile Phe Gly Leu Phe Glu Glu Thr Ala Leu Ala Ala Phe Leu Ser Tyr
945 950 955 960
Cys Pro Gly Met Asp Val Ala Leu Arg Met Tyr Pro Leu Lys Pro Ser
965 970 975
Trp Trp Phe Cys Ala Phe Pro Tyr Ser Phe Leu Ile Phe Val Tyr Asp
980 985 990
Glu Ile Arg Lys Leu Ile Leu Arg Arg Asn Pro Gly Gly Trp Val Glu
995 1000 1005
Lys Glu Thr Tyr Tyr
1010
<210> 7
<211> 1029
<212> PRT
<213> Homo sapiens
<400> 7
Met Gly Leu Trp Gly Lys Lys Gly Thr Val Ala Pro His Asp Gln Ser
1 5 10 15
Pro Arg Arg Arg Pro Lys Lys Gly Leu Ile Lys Lys Lys Met Val Lys
20 25 30
Arg Glu Lys Gln Lys Arg Asn Met Glu Glu Leu Lys Lys Glu Val Val
35 40 45
Met Asp Asp His Lys Leu Thr Leu Glu Glu Leu Ser Thr Lys Tyr Ser
50 55 60
Val Asp Leu Thr Lys Gly His Ser His Gln Arg Ala Lys Glu Ile Leu
65 70 75 80
Thr Arg Gly Gly Pro Asn Thr Val Thr Pro Pro Pro Thr Thr Pro Glu
85 90 95
Trp Val Lys Phe Cys Lys Gln Leu Phe Gly Gly Phe Ser Leu Leu Leu
100 105 110
Trp Thr Gly Ala Ile Leu Cys Phe Val Ala Tyr Ser Ile Gln Ile Tyr
115 120 125
Phe Asn Glu Glu Pro Thr Lys Asp Asn Leu Tyr Leu Ser Ile Val Leu
130 135 140
Ser Val Val Val Ile Val Thr Gly Cys Phe Ser Tyr Tyr Gln Glu Ala
145 150 155 160
Lys Ser Ser Lys Ile Met Glu Ser Phe Lys Asn Met Val Pro Gln Gln
165 170 175
Ala Leu Val Ile Arg Gly Gly Glu Lys Met Gln Ile Asn Val Gln Glu
180 185 190
Val Val Leu Gly Asp Leu Val Glu Ile Lys Gly Gly Asp Arg Val Pro
195 200 205
Ala Asp Leu Arg Leu Ile Ser Ala Gln Gly Cys Lys Val Asp Asn Ser
210 215 220
Ser Leu Thr Gly Glu Ser Glu Pro Gln Ser Arg Ser Pro Asp Phe Thr
225 230 235 240
His Glu Asn Pro Leu Glu Thr Arg Asn Ile Cys Phe Phe Ser Thr Asn
245 250 255
Cys Val Glu Gly Thr Ala Arg Gly Ile Val Ile Ala Thr Gly Asp Ser
260 265 270
Thr Val Met Gly Arg Ile Ala Ser Leu Thr Ser Gly Leu Ala Val Gly
275 280 285
Gln Thr Pro Ile Ala Ala Glu Ile Glu His Phe Ile His Leu Ile Thr
290 295 300
Val Val Ala Val Phe Leu Gly Val Thr Phe Phe Ala Leu Ser Leu Leu
305 310 315 320
Leu Gly Tyr Gly Trp Leu Glu Ala Ile Ile Phe Leu Ile Gly Ile Ile
325 330 335
Val Ala Asn Val Pro Glu Gly Leu Leu Ala Thr Val Thr Val Cys Leu
340 345 350
Thr Leu Thr Ala Lys Arg Met Ala Arg Lys Asn Cys Leu Val Lys Asn
355 360 365
Leu Glu Ala Val Glu Thr Leu Gly Ser Thr Ser Thr Ile Cys Ser Asp
370 375 380
Lys Thr Gly Thr Leu Thr Gln Asn Arg Met Thr Val Ala His Met Trp
385 390 395 400
Phe Asp Met Thr Val Tyr Glu Ala Asp Thr Thr Glu Glu Gln Thr Gly
405 410 415
Lys Thr Phe Thr Lys Ser Ser Asp Thr Trp Phe Met Leu Ala Arg Ile
420 425 430
Ala Gly Leu Cys Asn Arg Ala Asp Phe Lys Ala Asn Gln Glu Ile Leu
435 440 445
Pro Ile Ala Lys Arg Ala Thr Thr Gly Asp Ala Ser Glu Ser Ala Leu
450 455 460
Leu Lys Phe Ile Glu Gln Ser Tyr Ser Ser Val Ala Glu Met Arg Glu
465 470 475 480
Lys Asn Pro Lys Val Ala Glu Ile Pro Phe Asn Ser Thr Asn Lys Tyr
485 490 495
Gln Met Ser Ile His Leu Arg Glu Asp Ser Ser Gln Thr His Val Leu
500 505 510
Met Met Lys Gly Ala Pro Glu Arg Ile Leu Glu Phe Cys Ser Thr Phe
515 520 525
Leu Leu Asn Gly Gln Glu Tyr Ser Met Asn Asp Glu Met Lys Glu Ala
530 535 540
Phe Gln Asn Ala Tyr Leu Glu Leu Gly Gly Leu Gly Glu Arg Val Leu
545 550 555 560
Gly Phe Cys Phe Leu Asn Leu Pro Ser Ser Phe Ser Lys Gly Phe Pro
565 570 575
Phe Asn Thr Asp Glu Ile Asn Phe Pro Met Asp Asn Leu Cys Phe Val
580 585 590
Gly Leu Ile Ser Met Ile Asp Pro Pro Arg Ala Ala Val Pro Asp Ala
595 600 605
Val Ser Lys Cys Arg Ser Ala Gly Ile Lys Val Ile Met Val Thr Gly
610 615 620
Asp His Pro Ile Thr Ala Lys Ala Ile Ala Lys Gly Val Gly Ile Ile
625 630 635 640
Ser Glu Gly Thr Glu Thr Ala Glu Glu Val Ala Ala Arg Leu Lys Ile
645 650 655
Pro Ile Ser Lys Val Asp Ala Ser Ala Ala Lys Ala Ile Val Val His
660 665 670
Gly Ala Glu Leu Lys Asp Ile Gln Ser Lys Gln Leu Asp Gln Ile Leu
675 680 685
Gln Asn His Pro Glu Ile Val Phe Ala Arg Thr Ser Pro Gln Gln Lys
690 695 700
Leu Ile Ile Val Glu Gly Cys Gln Arg Leu Gly Ala Val Val Ala Val
705 710 715 720
Thr Gly Asp Gly Val Asn Asp Ser Pro Ala Leu Lys Lys Ala Asp Ile
725 730 735
Gly Ile Ala Met Gly Ile Ser Gly Ser Asp Val Ser Lys Gln Ala Ala
740 745 750
Asp Met Ile Leu Leu Asp Asp Asn Phe Ala Ser Ile Val Thr Gly Val
755 760 765
Glu Glu Gly Arg Leu Ile Phe Asp Asn Leu Lys Lys Ser Ile Met Tyr
770 775 780
Thr Leu Thr Ser Asn Ile Pro Glu Ile Thr Pro Phe Leu Met Phe Ile
785 790 795 800
Ile Leu Gly Ile Pro Leu Pro Leu Gly Thr Ile Thr Ile Leu Cys Ile
805 810 815
Asp Leu Gly Thr Asp Met Val Pro Ala Ile Ser Leu Ala Tyr Glu Ser
820 825 830
Ala Glu Ser Asp Ile Met Lys Arg Leu Pro Arg Asn Pro Lys Thr Asp
835 840 845
Asn Leu Val Asn His Arg Leu Ile Gly Met Ala Tyr Gly Gln Ile Gly
850 855 860
Met Ile Gln Ala Leu Ala Gly Phe Phe Thr Tyr Phe Val Ile Leu Ala
865 870 875 880
Glu Asn Gly Phe Arg Pro Val Asp Leu Leu Gly Ile Arg Leu His Trp
885 890 895
Glu Asp Lys Tyr Leu Asn Asp Leu Glu Asp Ser Tyr Gly Gln Gln Trp
900 905 910
Thr Tyr Glu Gln Arg Lys Val Val Glu Phe Thr Cys Gln Thr Ala Phe
915 920 925
Phe Val Thr Ile Val Val Val Gln Trp Ala Asp Leu Ile Ile Ser Lys
930 935 940
Thr Arg Arg Asn Ser Leu Phe Gln Gln Gly Met Arg Asn Lys Val Leu
945 950 955 960
Ile Phe Gly Ile Leu Glu Glu Thr Leu Leu Ala Ala Phe Leu Ser Tyr
965 970 975
Thr Pro Gly Met Asp Val Ala Leu Arg Met Tyr Pro Leu Lys Ile Thr
980 985 990
Trp Trp Leu Cys Ala Ile Pro Tyr Ser Ile Leu Ile Phe Val Tyr Asp
995 1000 1005
Glu Ile Arg Lys Leu Leu Ile Arg Gln His Pro Asp Gly Trp Val
1010 1015 1020
Glu Arg Glu Thr Tyr Tyr
1025
<210> 8
<211> 382
<212> PRT
<213> Homo sapiens
<400> 8
Met Gly Asp Trp Ser Ala Leu Gly Lys Leu Leu Asp Lys Val Gln Ala
1 5 10 15
Tyr Ser Thr Ala Gly Gly Lys Val Trp Leu Ser Val Leu Phe Ile Phe
20 25 30
Arg Ile Leu Leu Leu Gly Thr Ala Val Glu Ser Ala Trp Gly Asp Glu
35 40 45
Gln Ser Ala Phe Arg Cys Asn Thr Gln Gln Pro Gly Cys Glu Asn Val
50 55 60
Cys Tyr Asp Lys Ser Phe Pro Ile Ser His Val Arg Phe Trp Val Leu
65 70 75 80
Gln Ile Ile Phe Val Ser Val Pro Thr Leu Leu Tyr Leu Ala His Val
85 90 95
Phe Tyr Val Met Arg Lys Glu Glu Lys Leu Asn Lys Lys Glu Glu Glu
100 105 110
Leu Lys Val Ala Gln Thr Asp Gly Val Asn Val Asp Met His Leu Lys
115 120 125
Gln Ile Glu Ile Lys Lys Phe Lys Tyr Gly Ile Glu Glu His Gly Lys
130 135 140
Val Lys Met Arg Gly Gly Leu Leu Arg Thr Tyr Ile Ile Ser Ile Leu
145 150 155 160
Phe Lys Ser Ile Phe Glu Val Ala Phe Leu Leu Ile Gln Trp Tyr Ile
165 170 175
Tyr Gly Phe Ser Leu Ser Ala Val Tyr Thr Cys Lys Arg Asp Pro Cys
180 185 190
Pro His Gln Val Asp Cys Phe Leu Ser Arg Pro Thr Glu Lys Thr Ile
195 200 205
Phe Ile Ile Phe Met Leu Val Val Ser Leu Val Ser Leu Ala Leu Asn
210 215 220
Ile Ile Glu Leu Phe Tyr Val Phe Phe Lys Gly Val Lys Asp Arg Val
225 230 235 240
Lys Gly Lys Ser Asp Pro Tyr His Ala Thr Ser Gly Ala Leu Ser Pro
245 250 255
Ala Lys Asp Cys Gly Ser Gln Lys Tyr Ala Tyr Phe Asn Gly Cys Ser
260 265 270
Ser Pro Thr Ala Pro Leu Ser Pro Met Ser Pro Pro Gly Tyr Lys Leu
275 280 285
Val Thr Gly Asp Arg Asn Asn Ser Ser Cys Arg Asn Tyr Asn Lys Gln
290 295 300
Ala Ser Glu Gln Asn Trp Ala Asn Tyr Ser Ala Glu Gln Asn Arg Met
305 310 315 320
Gly Gln Ala Gly Ser Thr Ile Ser Asn Ser His Ala Gln Pro Phe Asp
325 330 335
Phe Pro Asp Asp Asn Gln Asn Ser Lys Lys Leu Ala Ala Gly His Glu
340 345 350
Leu Gln Pro Leu Ala Ile Val Asp Gln Arg Pro Ser Ser Arg Ala Ser
355 360 365
Ser Arg Ala Ser Ser Arg Pro Arg Pro Asp Asp Leu Glu Ile
370 375 380
<210> 9
<211> 354
<212> PRT
<213> Homo sapiens
<400> 9
Met Thr Ala Leu Ser Arg Ser Glu Ala Thr Glu Glu Gly Gly Asn Gln
1 5 10 15
Gln Met His Arg Lys Thr Ala Ser Leu Asn Ser Pro Val Ser Cys Lys
20 25 30
Glu Lys Pro Asp Arg Val Glu Glu Pro Pro Asp Tyr Ser Leu His Trp
35 40 45
Pro Glu Gly Leu Lys Gly Glu Glu Ile Lys Lys Cys Gly Arg Glu Gly
50 55 60
Ile Thr Leu Asn Lys Tyr Asn Gln Gln Tyr His Lys Leu Phe Lys Asp
65 70 75 80
Val Pro Leu Glu Glu Val Val Leu Lys Val Cys Ser Cys Ala Leu Gln
85 90 95
Arg Asp Phe Leu Leu Gln Gly Arg Leu Tyr Ile Ser Pro Asn Trp Leu
100 105 110
Cys Phe His Ala Ser Leu Phe Gly Lys Asp Ile Lys Val Val Ile Pro
115 120 125
Val Val Ser Val Gln Met Ile Lys Lys His Lys Met Ala Arg Leu Leu
130 135 140
Pro Asn Gly Leu Ala Ile Thr Thr Asn Thr Ser Gln Lys Tyr Ile Phe
145 150 155 160
Val Ser Leu Leu Ser Arg Asp Ser Val Tyr Asp Leu Leu Arg Arg Val
165 170 175
Cys Thr His Leu Gln Pro Ser Ser Lys Lys Ser Leu Ser Val Arg Glu
180 185 190
Phe Ser Gly Glu Pro Glu Ser Leu Glu Val Leu Ile Pro Glu Met Lys
195 200 205
Trp Arg Lys Val Cys Pro Ser Ser Arg Ser Leu Ser Leu Pro Asp Asn
210 215 220
Ile Pro Cys Ile Pro Pro Ser Ser Val Asp Ser Thr Asp Ser Phe Phe
225 230 235 240
Pro Ser Arg Lys Pro Pro Met Ser Glu Lys Ser Arg Ala Gln Val Ala
245 250 255
Ser Glu Asn Gly Gly Arg Trp Ala Trp Pro Met Pro Gly Trp Gly Pro
260 265 270
Ala Cys Pro Lys Lys Met Pro Asn Cys Ser Pro Thr Ala Lys Asn Ala
275 280 285
Val Tyr Glu Glu Asp Glu Leu Glu Glu Glu Pro Arg Ser Thr Gly Glu
290 295 300
Leu Arg Leu Trp Asp Tyr Arg Leu Leu Lys Val Phe Phe Val Leu Ile
305 310 315 320
Cys Phe Leu Val Met Ser Ser Ser Tyr Leu Ala Phe Arg Ile Ser Arg
325 330 335
Leu Glu Gln Gln Leu Cys Ser Leu Ser Trp Asp Asp Pro Val Pro Gly
340 345 350
His Arg
<210> 10
<211> 432
<212> PRT
<213> Homo sapiens
<400> 10
Met Thr Glu Leu Gln Gln Asp Val Glu Asp Thr Lys Pro Ala Lys Val
1 5 10 15
Leu Gly Lys Arg Glu Ser Lys Leu Gly Ser Ala His Ser Glu Ala Glu
20 25 30
Asn Gly Val Glu Glu Lys Lys Lys Ala Cys Arg Ser Pro Thr Ala Gln
35 40 45
Ser Pro Thr Pro Ser Val Glu Ala Asp Ser Pro Asp Gln Lys Lys Ile
50 55 60
Ile Ser Leu Trp Ser Lys Ser Ser Phe Asp Gly Ala Ser Leu Ala Ser
65 70 75 80
Asp Lys Asn Asp Cys Lys Thr Glu Ser Lys Asn Asp Pro Lys Thr Glu
85 90 95
Arg Lys Lys Ser Ser Ser Ser Ser Gln Tyr Lys Ala Asn Met His Phe
100 105 110
His Lys Leu Phe Leu Ser Val Pro Thr Glu Glu Pro Leu Lys Gln Ser
115 120 125
Phe Thr Cys Ala Leu Gln Lys Glu Ile Leu Tyr Gln Gly Lys Leu Phe
130 135 140
Val Ser Glu Asn Trp Ile Cys Phe His Ser Lys Val Phe Gly Lys Asp
145 150 155 160
Thr Lys Ile Ser Ile Pro Ala Phe Ser Val Thr Leu Ile Lys Lys Thr
165 170 175
Lys Thr Ala Leu Leu Val Pro Asn Ala Leu Ile Ile Ala Thr Val Thr
180 185 190
Asp Arg Tyr Ile Phe Val Ser Leu Leu Ser Arg Asp Ser Thr Tyr Lys
195 200 205
Leu Leu Lys Ser Val Cys Gly His Leu Glu Asn Thr Ser Val Gly Asn
210 215 220
Ser Pro Asn Pro Ser Ser Ala Glu Asn Ser Phe Arg Ala Asp Arg Pro
225 230 235 240
Ser Ser Leu Pro Leu Asp Phe Asn Asp Glu Phe Ser Asp Leu Asp Gly
245 250 255
Val Val Gln Gln Arg Arg Gln Asp Met Glu Gly Tyr Ser Ser Ser Gly
260 265 270
Ser Gln Thr Pro Glu Ser Glu Asn Ser Arg Asp Phe His Ala Thr Glu
275 280 285
Ser Gln Thr Val Leu Asn Val Ser Lys Gly Glu Ala Lys Pro Thr Arg
290 295 300
Ala Asp Ala His Val Asn Arg Val Pro Glu Gly Lys Ala Lys Ser Leu
305 310 315 320
Pro Val Gln Gly Leu Ser Glu Thr Val Gly Ile Leu His Lys Val Lys
325 330 335
Ser Gln Lys Cys Pro Met Leu His His Ile Leu Ile Phe Tyr Ala Ile
340 345 350
Val Val Cys Ala Leu Ile Ile Ser Thr Phe Tyr Met Arg Tyr Arg Ile
355 360 365
Asn Thr Leu Glu Glu Gln Leu Gly Leu Leu Thr Ser Ile Val Asp Thr
370 375 380
His Asn Thr Glu Gln Ala Ala Pro Ser Gly Leu Arg Ser Gln Val Gln
385 390 395 400
Phe Asn Val Glu Val Leu Cys Gln Glu Leu Thr Ala Asn Ile Val Lys
405 410 415
Leu Glu Lys Ile Gln Asn Asn Leu Gln Lys Leu Leu Glu Asn Gly Asp
420 425 430
<210> 11
<211> 197
<212> PRT
<213> Homo sapiens
<400> 11
Met Asp Val Gly Ser Lys Glu Val Leu Met Glu Ser Pro Pro Asp Tyr
1 5 10 15
Ser Ala Ala Pro Arg Gly Arg Phe Gly Ile Pro Cys Cys Pro Val His
20 25 30
Leu Lys Arg Leu Leu Ile Val Val Val Val Val Val Leu Ile Val Val
35 40 45
Val Ile Val Gly Ala Leu Leu Met Gly Leu His Met Ser Gln Lys His
50 55 60
Thr Glu Met Val Leu Glu Met Ser Ile Gly Ala Pro Glu Ala Gln Gln
65 70 75 80
Arg Leu Ala Leu Ser Glu His Leu Val Thr Thr Ala Thr Phe Ser Ile
85 90 95
Gly Ser Thr Gly Leu Val Val Tyr Asp Tyr Gln Gln Leu Leu Ile Ala
100 105 110
Tyr Lys Pro Ala Pro Gly Thr Cys Cys Tyr Ile Met Lys Ile Ala Pro
115 120 125
Glu Ser Ile Pro Ser Leu Glu Ala Leu Asn Arg Lys Val His Asn Phe
130 135 140
Gln Met Glu Cys Ser Leu Gln Ala Lys Pro Ala Val Pro Thr Ser Lys
145 150 155 160
Leu Gly Gln Ala Glu Gly Arg Asp Ala Gly Ser Ala Pro Ser Gly Gly
165 170 175
Asp Pro Ala Phe Leu Gly Met Ala Val Asn Thr Leu Cys Gly Glu Val
180 185 190
Pro Leu Tyr Tyr Ile
195
<210> 12
<211> 562
<212> PRT
<213> Homo sapiens
<400> 12
Met Ala Ile Ile Tyr Leu Ile Leu Leu Phe Thr Ala Val Arg Gly Asp
1 5 10 15
Gln Ile Cys Ile Gly Tyr His Ala Asn Asn Ser Thr Glu Lys Val Asp
20 25 30
Thr Asn Leu Glu Arg Asn Val Thr Val Thr His Ala Lys Asp Ile Leu
35 40 45
Glu Lys Thr His Asn Gly Lys Leu Cys Lys Leu Asn Gly Ile Pro Pro
50 55 60
Leu Glu Leu Gly Asp Cys Ser Ile Ala Gly Trp Leu Leu Gly Asn Pro
65 70 75 80
Glu Cys Asp Arg Leu Leu Ser Val Pro Glu Trp Ser Tyr Ile Met Glu
85 90 95
Lys Glu Asn Pro Arg Asp Gly Leu Cys Tyr Pro Gly Ser Phe Asn Asp
100 105 110
Tyr Glu Glu Leu Lys His Leu Leu Ser Ser Val Lys His Phe Glu Lys
115 120 125
Val Lys Ile Leu Pro Lys Asp Arg Trp Thr Gln His Thr Thr Thr Gly
130 135 140
Gly Ser Arg Ala Cys Ala Val Ser Gly Asn Pro Ser Phe Phe Arg Asn
145 150 155 160
Met Val Trp Leu Thr Lys Glu Gly Ser Asp Tyr Pro Val Ala Lys Gly
165 170 175
Ser Tyr Asn Asn Thr Ser Gly Glu Gln Met Leu Ile Ile Trp Gly Val
180 185 190
His His Pro Ile Asp Glu Thr Glu Gln Arg Thr Leu Tyr Gln Asn Val
195 200 205
Gly Thr Tyr Val Ser Val Gly Thr Ser Thr Leu Asn Lys Arg Ser Thr
210 215 220
Pro Glu Ile Ala Thr Arg Pro Lys Val Asn Gly Gln Gly Gly Arg Met
225 230 235 240
Glu Phe Ser Trp Thr Leu Leu Asp Met Trp Asp Thr Ile Asn Phe Glu
245 250 255
Ser Thr Gly Asn Leu Ile Ala Pro Glu Tyr Gly Phe Lys Ile Ser Lys
260 265 270
Arg Gly Ser Ser Gly Ile Met Lys Thr Glu Gly Thr Leu Glu Asn Cys
275 280 285
Glu Thr Lys Cys Gln Thr Pro Leu Gly Ala Ile Asn Thr Thr Leu Pro
290 295 300
Phe His Asn Val His Pro Leu Thr Ile Gly Glu Cys Pro Lys Tyr Val
305 310 315 320
Lys Ser Glu Lys Leu Val Leu Ala Thr Gly Leu Arg Asn Val Pro Gln
325 330 335
Ile Glu Ser Arg Gly Leu Phe Gly Ala Ile Ala Gly Phe Ile Glu Gly
340 345 350
Gly Trp Gln Gly Met Val Asp Gly Trp Tyr Gly Tyr His His Ser Asn
355 360 365
Asp Gln Gly Ser Gly Tyr Ala Ala Asp Lys Glu Ser Thr Gln Lys Ala
370 375 380
Phe Asp Gly Ile Thr Asn Lys Val Asn Ser Val Ile Glu Lys Met Asn
385 390 395 400
Thr Gln Phe Glu Ala Val Gly Lys Glu Phe Gly Asn Leu Glu Arg Arg
405 410 415
Leu Glu Asn Leu Asn Lys Arg Met Glu Asp Gly Phe Leu Asp Val Trp
420 425 430
Thr Tyr Asn Ala Glu Leu Leu Val Leu Met Glu Asn Glu Arg Thr Leu
435 440 445
Asp Phe His Asp Ser Asn Val Lys Asn Leu Tyr Asp Lys Val Arg Met
450 455 460
Gln Leu Arg Asp Asn Val Lys Glu Leu Gly Asn Gly Cys Phe Glu Phe
465 470 475 480
Tyr His Lys Cys Asp Asp Glu Cys Met Asn Ser Val Lys Asn Gly Thr
485 490 495
Tyr Asp Tyr Pro Lys Tyr Glu Glu Glu Ser Lys Leu Asn Arg Asn Glu
500 505 510
Ile Lys Gly Val Lys Leu Ser Ser Met Gly Val Tyr Gln Ile Leu Ala
515 520 525
Ile Tyr Ala Thr Val Ala Gly Ser Leu Ser Leu Ala Ile Met Met Ala
530 535 540
Gly Ile Ser Phe Trp Met Cys Ser Asn Gly Ser Leu Gln Cys Arg Ile
545 550 555 560
Cys Ile
<210> 13
<211> 566
<212> PRT
<213> Homo sapiens
<400> 13
Met Lys Thr Ile Ile Ala Leu Ser Tyr Ile Phe Cys Leu Ala Leu Gly
1 5 10 15
Gln Asp Leu Pro Gly Asn Asp Asn Asn Thr Ala Thr Leu Cys Leu Gly
20 25 30
His His Ala Val Pro Asn Gly Thr Leu Val Lys Thr Ile Thr Asp Asp
35 40 45
Gln Ile Glu Val Thr Asn Ala Thr Glu Leu Val Gln Ser Ser Ser Thr
50 55 60
Gly Lys Ile Cys Asn Asn Pro His Arg Ile Leu Asp Gly Ile Asp Cys
65 70 75 80
Thr Leu Ile Asp Ala Leu Leu Gly Asp Pro His Cys Asp Val Phe Gln
85 90 95
Asn Glu Thr Trp Asp Leu Phe Val Glu Arg Ser Lys Ala Phe Ser Asn
100 105 110
Cys Tyr Pro Tyr Asp Val Pro Asp Tyr Ala Ser Leu Arg Ser Leu Val
115 120 125
Ala Ser Ser Gly Thr Leu Glu Phe Ile Thr Glu Gly Phe Thr Trp Thr
130 135 140
Gly Val Thr Gln Asn Gly Gly Ser Asn Ala Cys Lys Arg Gly Pro Asp
145 150 155 160
Ser Gly Phe Phe Ser Arg Leu Asn Trp Leu Thr Lys Ser Gly Ser Thr
165 170 175
Tyr Pro Val Leu Asn Val Thr Met Pro Asn Asn Asp Asn Phe Asp Lys
180 185 190
Leu Tyr Ile Trp Gly Val His His Pro Ser Thr Asn Gln Glu Gln Thr
195 200 205
Ser Leu Tyr Val Gln Ala Ser Gly Arg Val Thr Val Ser Thr Arg Arg
210 215 220
Ser Gln Gln Thr Ile Ile Pro Asn Ile Gly Ser Arg Pro Trp Val Arg
225 230 235 240
Gly Gln Ser Ser Arg Ile Ser Ile Tyr Trp Thr Ile Val Lys Pro Gly
245 250 255
Asp Val Leu Val Ile Asn Ser Asn Gly Asn Leu Ile Ala Pro Arg Gly
260 265 270
Tyr Phe Lys Met Arg Thr Gly Lys Ser Ser Ile Met Arg Ser Asp Ala
275 280 285
Pro Ile Asp Thr Cys Ile Ser Glu Cys Ile Thr Pro Asn Gly Ser Ile
290 295 300
Pro Asn Asp Lys Pro Phe Gln Asn Val Asn Lys Ile Thr Tyr Gly Ala
305 310 315 320
Cys Pro Lys Tyr Val Lys Gln Asn Thr Leu Lys Leu Ala Thr Gly Met
325 330 335
Arg Asn Val Pro Glu Lys Gln Thr Arg Gly Leu Phe Gly Ala Ile Ala
340 345 350
Gly Phe Ile Glu Asn Gly Trp Glu Gly Met Ile Asp Gly Trp Tyr Gly
355 360 365
Phe Arg His Gln Asn Ser Glu Gly Thr Gly Gln Ala Ala Asp Leu Lys
370 375 380
Ser Thr Gln Ala Ala Ile Asp Gln Ile Asn Gly Lys Leu Asn Arg Val
385 390 395 400
Ile Glu Lys Thr Asn Glu Lys Phe His Gln Ile Glu Lys Glu Phe Ser
405 410 415
Glu Val Glu Gly Arg Ile Gln Asp Leu Glu Lys Tyr Val Glu Asp Thr
420 425 430
Lys Ile Asp Leu Trp Ser Tyr Asn Ala Glu Leu Leu Val Ala Leu Glu
435 440 445
Asn Gln His Thr Ile Asp Leu Thr Asp Ser Glu Met Asn Lys Leu Phe
450 455 460
Glu Lys Thr Arg Arg Gln Leu Arg Glu Asn Ala Glu Asp Met Gly Asn
465 470 475 480
Gly Cys Phe Lys Ile Tyr His Lys Cys Asp Asn Ala Cys Ile Glu Ser
485 490 495
Ile Arg Asn Gly Thr Tyr Asp His Asp Val Tyr Arg Asp Glu Ala Leu
500 505 510
Asn Asn Arg Phe Gln Ile Lys Gly Val Glu Leu Lys Ser Gly Tyr Lys
515 520 525
Asp Trp Ile Leu Trp Ile Ser Phe Ala Ile Ser Cys Phe Leu Leu Cys
530 535 540
Val Val Leu Leu Gly Phe Ile Met Trp Ala Cys Gln Arg Gly Asn Ile
545 550 555 560
Arg Cys Asn Ile Cys Ile
565
<210> 14
<211> 469
<212> PRT
<213> Homo sapiens
<400> 14
Met Asn Pro Asn Gln Lys Ile Ile Thr Ile Gly Ser Val Ser Leu Thr
1 5 10 15
Ile Ala Thr Val Cys Phe Leu Met Gln Ile Ala Ile Leu Ala Thr Thr
20 25 30
Val Thr Leu His Phe Lys Gln His Lys Cys Asp Ser Pro Ala Ser Asn
35 40 45
Gln Val Met Pro Cys Glu Pro Ile Ile Ile Glu Arg Asn Ile Thr Glu
50 55 60
Ile Val Tyr Leu Asn Asn Thr Thr Ile Glu Lys Glu Ile Cys Pro Glu
65 70 75 80
Val Val Glu Tyr Arg Asn Trp Ser Lys Pro Gln Cys Gln Ile Thr Gly
85 90 95
Phe Ala Pro Phe Ser Lys Asp Asn Ser Ile Arg Leu Ser Ala Gly Gly
100 105 110
Asp Ile Trp Val Thr Arg Glu Pro Tyr Val Ser Cys Asp Pro Gly Lys
115 120 125
Cys Tyr Gln Phe Ala Leu Gly Gln Gly Thr Thr Leu Asp Asn Lys His
130 135 140
Ser Asn Gly Thr Ile His Asp Arg Ile Pro His Arg Thr Leu Leu Met
145 150 155 160
Asn Glu Leu Gly Val Pro Phe His Leu Gly Thr Lys Gln Val Cys Val
165 170 175
Ala Trp Ser Ser Ser Ser Cys His Asp Gly Lys Ala Trp Leu His Val
180 185 190
Cys Val Thr Gly Asp Asp Arg Asn Ala Thr Ala Ser Phe Ile Tyr Asp
195 200 205
Gly Arg Leu Val Asp Ser Ile Gly Ser Trp Ser Gln Asn Ile Leu Arg
210 215 220
Thr Gln Glu Ser Glu Cys Val Cys Ile Asn Gly Thr Cys Thr Val Val
225 230 235 240
Met Thr Asp Gly Ser Ala Ser Gly Arg Ala Asp Thr Arg Ile Leu Phe
245 250 255
Ile Lys Glu Gly Lys Ile Val His Ile Gly Pro Leu Ser Gly Ser Ala
260 265 270
Gln His Ile Glu Glu Cys Ser Cys Tyr Pro Arg Tyr Pro Asp Val Arg
275 280 285
Cys Ile Cys Arg Asp Asn Trp Lys Gly Ser Asn Arg Pro Val Ile Asp
290 295 300
Ile Asn Met Glu Asp Tyr Ser Ile Asp Ser Ser Tyr Val Cys Ser Gly
305 310 315 320
Leu Val Gly Asp Thr Pro Arg Asn Asp Asp Ser Ser Ser Asn Ser Asn
325 330 335
Cys Arg Asp Pro Asn Asn Glu Arg Gly Asn Pro Gly Val Lys Gly Trp
340 345 350
Ala Phe Asp Asn Gly Asp Asp Val Trp Met Gly Arg Thr Ile Ser Lys
355 360 365
Asp Leu Arg Ser Gly Tyr Glu Thr Phe Lys Val Ile Gly Gly Trp Ser
370 375 380
Thr Pro Asn Ser Lys Ser Gln Val Asn Arg Gln Val Ile Val Asp Asn
385 390 395 400
Asn Asn Trp Ser Gly Tyr Ser Gly Ile Phe Ser Val Glu Gly Lys Ser
405 410 415
Cys Ile Asn Arg Cys Phe Tyr Val Glu Leu Ile Arg Gly Arg Pro Gln
420 425 430
Glu Thr Arg Val Trp Trp Thr Ser Asn Ser Ile Val Val Phe Cys Gly
435 440 445
Thr Ser Gly Thr Tyr Gly Thr Gly Ser Trp Pro Asp Gly Ala Asn Ile
450 455 460
Asn Phe Met Pro Ile
465
<210> 15
<211> 469
<212> PRT
<213> Homo sapiens
<400> 15
Met Asn Pro Asn Gln Lys Ile Ile Thr Ile Gly Ser Val Ser Leu Thr
1 5 10 15
Ile Ala Thr Val Cys Phe Leu Met Gln Thr Ala Ile Leu Val Thr Thr
20 25 30
Val Thr Leu His Phe Lys Gln Tyr Glu Cys Asp Ser Pro Ala Ser Asn
35 40 45
Gln Val Met Pro Cys Glu Pro Ile Ile Ile Glu Arg Asn Ile Thr Glu
50 55 60
Ile Val Tyr Leu Asn Asn Thr Thr Ile Glu Lys Glu Ile Cys Pro Lys
65 70 75 80
Val Val Glu Tyr Arg Asn Trp Ser Lys Pro Gln Cys Gln Ile Thr Gly
85 90 95
Phe Ala Pro Phe Ser Lys Asp Asn Ser Ile Arg Leu Ser Ala Gly Gly
100 105 110
Asp Ile Trp Val Thr Arg Glu Pro Tyr Val Ser Cys Asp His Gly Lys
115 120 125
Cys Tyr Gln Phe Ala Leu Gly Gln Gly Thr Thr Leu Asp Asn Lys His
130 135 140
Ser Asn Asp Thr Ile His Asp Arg Ile Pro His Arg Thr Leu Leu Met
145 150 155 160
Asn Glu Leu Gly Val Pro Phe His Leu Gly Thr Arg Gln Val Cys Ile
165 170 175
Ala Trp Ser Ser Ser Ser Cys His Asp Gly Lys Ala Trp Leu His Val
180 185 190
Cys Ile Thr Gly Asp Asp Lys Asn Ala Thr Ala Ser Phe Ile Tyr Asp
195 200 205
Gly Arg Leu Val Asp Ser Ile Gly Ser Trp Ser Gln Asn Ile Leu Arg
210 215 220
Thr Gln Glu Ser Glu Cys Val Cys Ile Asn Gly Thr Cys Thr Val Val
225 230 235 240
Met Thr Asp Gly Ser Ala Ser Gly Arg Ala Asp Thr Arg Ile Leu Phe
245 250 255
Ile Glu Glu Gly Lys Ile Val His Ile Ser Pro Leu Ser Gly Ser Ala
260 265 270
Gln His Val Glu Glu Cys Ser Cys Tyr Pro Arg Tyr Pro Gly Val Arg
275 280 285
Cys Ile Cys Arg Asp Asn Trp Lys Gly Ser Asn Arg Pro Val Val Asp
290 295 300
Ile Asn Met Glu Asp Tyr Ser Ile Asp Ser Ser Tyr Val Cys Ser Gly
305 310 315 320
Leu Val Gly Asp Thr Pro Arg Asn Asp Asp Arg Ser Ser Asn Ser Asn
325 330 335
Cys Arg Asn Pro Asn Asn Glu Arg Gly Asn Gln Gly Val Lys Gly Trp
340 345 350
Ala Phe Asp Asn Gly Asp Asp Val Trp Met Gly Arg Thr Ile Ser Lys
355 360 365
Asp Leu Arg Ser Gly Tyr Glu Thr Phe Lys Val Ile Gly Gly Trp Ser
370 375 380
Thr Pro Asn Ser Lys Ser Gln Ile Asn Arg Gln Val Ile Val Asp Ser
385 390 395 400
Asp Asn Arg Ser Gly Tyr Ser Gly Ile Phe Ser Val Glu Gly Lys Ser
405 410 415
Cys Ile Asn Arg Cys Phe Tyr Val Glu Leu Ile Arg Gly Arg Lys Gln
420 425 430
Glu Ala Arg Val Trp Trp Thr Ser Asn Ser Ile Val Val Phe Cys Gly
435 440 445
Thr Ser Gly Thr Tyr Gly Thr Gly Ser Trp Pro Asp Gly Ala Asn Ile
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Asn Phe Met Pro Ile
465
Claims (15)
- i) 폐 상피 세포에서 정단(apical) 나트륨-칼륨-ATPase(sodium-potassium-ATPase; NKA) 국소화를 방지하거나,
ii) ROCK 억제제의 투여 전 정단 NKA 국소화와 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시킴으로써,
폐부종의 치료 또는 예방에서의 용도를 위한 ROCK 억제제로서,
폐부종은 바이러스 감염과 관련이 있고,
바이러스는 인플루엔자 바이러스인, ROCK 억제제. - 제1항에 있어서, ROCK 억제제는 ROCK 1 또는 ROCK 1/2 억제제인, ROCK 억제제.
- 제1항에 있어서, ROCK 억제제는 파수딜(Fasudil), Rho XIII, Y27632, 하이드록시파수딜(Hydroxyfasudil), H-1152-P, Y27632, Y30141, Y32885, Y39983, DW1865, SLx-2119, SR8046, SR6246, 리파수딜(Ripasudil), AS1892892, AR12141, AR12432, INS-117548, INS-115644, AT13148, RKI1447, SAR407899, 네타르수딜(Netarsudil), AR12286, PG286*, PG324**, ATS907, AMA0076, 티아조비빈(Thiazovivin), 아자벤즈이미다졸-아미노푸라잔(Azabenzimidazole-aminofurazans), H-0104 디하이드로클로라이드(Dihydrochloride), DE-104, 올레핀, 이소퀴놀린, 인다졸, 피리딘알켄 유도체, H-1152 디클로라이드(dichloride), XD-4000, HMN-1152, 4-(1-아미노알킬)-N-(4-피리딜)사이클로헥산-카르복사미드, 로스타틴(Rhostatin), BA-210, BA-207, BA-215, BA-285, BA-1037, Ki-23095, VAS-012, 퀴나졸린, AR13154 및/또는 AMA0428 또는 이들의 조합으로 이루어진 군으로부터 선택되는, ROCK 억제제.
- 제1항 내지 제3항 중 어느 한 항에 있어서, 폐부종은 흉부 엑스레이 또는 컴퓨터 단층촬영에 의해 진단되는, ROCK 억제제.
- 제1항 내지 제4항 중 어느 한 항에 있어서, ROCK 억제제는 인플루엔자 A 또는 인플루엔자 B 바이러스에 감염되거나 감염 위험이 있는 대상체에게 투여되는, ROCK 억제제.
- 제1항 내지 제5항 중 어느 한 항에 있어서, 폐 상피 세포는 폐포 상피 세포 및/또는 기관지 상피 세포인, ROCK 억제제.
- 제1항 내지 제6항 중 어느 한 항에 있어서, 폐포 상피 세포는 타입 I 또는 타입 II 폐포 상피 세포인, ROCK 억제제.
- 제1항 내지 제7항 중 어느 한 항에 있어서, 기관지 상피 세포는 섬모 및/또는 비섬모 기관지 상피 세포인, ROCK 억제제.
- 제1항 내지 제8항 중 어느 한 항에 있어서, ROCK 억제제는 시험관내 시험 시스템과 접촉할 때, 접촉 전 시험관내 시험 시스템에서 폐 상피 세포의 정단 NKA 국소화와 비교하여, 폐 상피 세포에서 정단 NKA 국소화를 감소시키고, 시험 시스템은 인플루엔자 바이러스로 감염되는 배양된 폐 상피 세포를 포함하는, ROCK 억제제.
- 제1항 내지 제9항 중 어느 한 항에 있어서, ROCK 억제제는 ROCK 억제제의 투여 전 바이러스 부하(virus load)와 비교하여 바이러스 부하를 감소시키는, ROCK 억제제.
- 제1항 내지 제10항 중 어느 한 항에 있어서, ROCK 억제제는 ROCK 억제제의 투여 전에 존재하는 폐의 체액 중량과 비교하여 폐의 체액 중량을 감소시키는, ROCK 억제제.
- 제1항 내지 제11항 중 어느 한 항에 있어서, ROCK 억제제는 ROCK 억제제의 투여 전에 대식세포의 폐로의 침윤(infiltration)과 비교하여 폐로의 대식세포의 침윤을 감소시키는, ROCK 억제제.
- 폐 상피 세포에서 정단 NKA 국소화를 방지하거나 감소시키는 데 효과적인 억제제의 결정을 위한 인플루엔자 바이러스로 감염되는 배양된 폐 상피 세포를 포함하는 시험관내(in vitro) 시험 시스템의 용도.
- 폐부종의 예방 및/또는 치료에 효과적인 분자를 검출하는 방법으로서, 관심 화합물과 인플루엔자 바이러스에 감염되는 배양된 폐 상피 세포를 포함하는 시험관내 시험 시스템을 접촉시키는 것을 포함하고, 관심 화합물은 접촉 전의 시험관내 시험 시스템과 비교하여 폐 상피 세포에서 정단 NKA 국소화를 감소시키는, 방법.
- i) ROCK 억제제;
ii) 폐 상피 세포;
iii) 인플루엔자 바이러스; 및
iv) NKA의 검출 및 세포 국소화를 위한 수단
을 포함하는, 시험 시스템.
Applications Claiming Priority (3)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| LULU101746 | 2020-04-22 | ||
| LU101746 | 2020-04-22 | ||
| PCT/EP2021/060490 WO2021214200A1 (en) | 2020-04-22 | 2021-04-22 | Rock inhibitors for use in treating or preventing pulmonary edema |
Publications (1)
| Publication Number | Publication Date |
|---|---|
| KR20230004706A true KR20230004706A (ko) | 2023-01-06 |
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| Application Number | Title | Priority Date | Filing Date |
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| KR1020227040455A KR20230004706A (ko) | 2020-04-22 | 2021-04-22 | 폐부종의 치료 또는 예방용 rock 억제제 |
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| Country | Link |
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| US (1) | US20230172944A1 (ko) |
| EP (1) | EP4138846A1 (ko) |
| JP (1) | JP2023523730A (ko) |
| KR (1) | KR20230004706A (ko) |
| CN (1) | CN115666580A (ko) |
| AU (1) | AU2021259332A1 (ko) |
| BR (1) | BR112022019955A2 (ko) |
| CA (1) | CA3173018A1 (ko) |
| MX (1) | MX2022013196A (ko) |
| WO (1) | WO2021214200A1 (ko) |
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| CA2781390A1 (en) * | 2009-12-14 | 2011-06-23 | Inspire Pharmaceuticals, Inc. | Bridged bicyclic rho kinase inhibitor compounds, composition and use |
| JP2013532689A (ja) | 2010-07-27 | 2013-08-19 | インスパイアー ファーマシューティカルズ，インコーポレイティド | プロドラッグ形態のキナーゼ阻害剤化合物を使用して眼疾患を治療する方法 |
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| US20230172944A1 (en) | 2023-06-08 |
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| CN115666580A (zh) | 2023-01-31 |
| AU2021259332A1 (en) | 2022-10-27 |
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| JP2023523730A (ja) | 2023-06-07 |
| WO2021214200A1 (en) | 2021-10-28 |
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