KR101685109B1 - 근육 분화, 재생, 또는 질환 상태와 관련된 노고-에이의 용도 - Google Patents
근육 분화, 재생, 또는 질환 상태와 관련된 노고-에이의 용도 Download PDFInfo
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- KR101685109B1 KR101685109B1 KR1020140186373A KR20140186373A KR101685109B1 KR 101685109 B1 KR101685109 B1 KR 101685109B1 KR 1020140186373 A KR1020140186373 A KR 1020140186373A KR 20140186373 A KR20140186373 A KR 20140186373A KR 101685109 B1 KR101685109 B1 KR 101685109B1
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Abstract
근육 분화, 재생, 또는 질환 상태와 관련된 노고-에이의 용도에 관한 것이다. 노고-에이는 근육의 분화 또는 재생 정도, 또는 질환 상태를 확인하기 위해 이용될 수 있다.
Description
근육 분화, 재생, 또는 질환 상태와 관련된 노고-에이의 용도에 관한 것이다.
노고-에이(Nogo-A)는 노고 관련 세 가지 이성질체 중 가장 길이가 긴 단백질이다. 이들 세 이성질체는 유전자 전사과정 중 프로모터 혹은 스플라이싱의 차이로 인해 만들어지고, 이들은 3` 말단에 잘 보전된 공통된 부분을 가진다. 이를 제외한 5` 말단 부분은 이성질체 간에 차이를 가지고 있고, 이로 인해 이들의 작용 효과와 발현 양상은 크게 상이하게 된다.
본 발명에서 다룬 노고-에이는 중추 신경계에서의 작용 효과가 기존 연구들에 의해 잘 규명되어 있다. 중추 신경계에서 노고-에이는 희소돌기아교세포의 말이집 형성을 억제하고, 뉴론 축삭의 길이 성장을 방해하며, 신경 조직의 재생 효과를 저해하는 작용을 한다.
그러나 노고-에이의 신경계 외에서의 작용 효과에 대해선 잘 알려져 있지 않다. 흔히 루게릭병으로 알려진 근육위축가쪽경화증은 운동신경세포의 선택적 사멸로 인해 전신의 근육이 쇄약해지고 위축되는 질병인데, 이 질병 상태에 있는 근육에서 병증에 따라 노고-에이의 발현이 일치하는 경향을 보인다. 이처럼 근육에서의 노고-에이의 작용 또한 신경계와 관련되어 설명하는 수준에 그치고 있다. 온전히 근육 자체에서의 노고-에이의 발현에 대해선 현재 진행된 연구가 전무하다.
이에 본 발명자는 노고-에이의 근육에서의 발현 양상에 대한 내용을 제공하고, 근육 분화 및 재생 그리고 질환 상태 표지 인자로서의 새로운 기능을 제공한다.
일 양상은 근육전구세포의 근육세포로의 분화 정도 확인용 조성물 또는 키트를 제공한다.
다른 양상은 근육전구세포의 근육세포로의 분화 정도를 확인하는 방법을 제공한다.
다른 양상은 손상된 근육의 재생 정도 확인용 조성물 또는 키트를 제공한다.
다른 양상은 손상된 근육의 재생 정도를 확인하는 방법을 제공한다.
다른 양상은 근육 손상 질환 진단용 조성물 또는 키트를 제공한다.
다른 양상은 개체의 근육 손상 질환을 진단하기 위한 정보제공방법을 제공한다.
일 양상은 노고-에이(Nogo-A) 유전자의 mRNA 또는 단백질의 발현 수준을 측정하는 제제를 포함하는, 근육전구세포의 근육세포로의 분화 정도 확인용 조성물을 제공한다.
상기 노고-에이는 신경돌기성장 억제제 A(neurite outgrowth inhibitor A)로서, 레티쿨론 4(reticulon 4) isoform A, 또는 RTN4-A로 명명될 수 있다. 상기 노고-에이 유전자는 서열번호 1의 폴리펩티드를 코딩하는 것일 수 있다. 상기 노고-에이 유전자는 서열번호 2의 폴리뉴클레오티드를 전사체로 포함하는 것일 수 있다.
근육전구세포(precursor cell)는 근육세포로 발달 또는 분화할 수 있는 세포를 의미한다. 근육전구세포는 근육 원조세포(progenitor cell) 및 근육 줄기세포를 포함한다. 근육전구세포는 위성세포(satellite cell) 또는 근모세포(myoblast)일 수 있다. 근육세포는 근세포(myocyte) 또는 근관세포(myotube)일 수 있다.
상기 제제는 상기 유전자에 특이적으로 결합하는 프라이머 쌍, 프로브, 또는 상기 단백질에 특이적으로 결합하는 항체일 수 있다. 용어 "프라이머(primer)"는 3' 말단에 자유 히드록실기를 갖고, 상보적인 서열을 갖는 주형과 염기쌍을 형성함으로써 주형 복제를 위한 시작점으로 기능하는 짧은 올리고뉴클레오티드를 의미한다. 노고-에이 유전자에 특이적으로 결합하는 프라이머 쌍의 각 서열, 중합효소연쇄반응 조건, 및 각 프라이머의 길이는 노고-에이 유전자의 뉴클레오티드 서열을 이용하여 통상의 기술자가 적절하게 설계 또는 변형시킬 수 있다.
용어 "프로브(probe)"는 시료 중 그의 서열과 상보적인 서열을 갖는 표적 뉴클레오티드의 존재를 검출할 수 있는 핵산 단편을 의미한다. 표지된 프로브를 단일가닥 핵산으로 변성시킨 후 표적 핵산과의 혼성화를 유도함으로써 표적 뉴클레오티드의 존재를 검출할 수 있다. 노고-에이 유전자에 특이적으로 결합하는 프로브 및 그의 표적 서열과의 혼성화 조건은 통상의 기술자가 적절하게 설계 또는 변형시킬 수 있다.
용어 "항체(antibody)"는 항원의 특정 부위를 인식하는 특이적인 단백질 분자를 의미한다. 노고-에이 단백질에 특이적으로 결합하는 항체는 상기 단백질에 특이성 및 친화성이 높고 다른 단백질에 대한 교차 반응성이 거의 없는 항체로, 단클론 항체, 다클론 항체 또는 재조합 항체일 수 있다.
다른 양상은 근육전구세포의 근육세포로의 분화 정도 확인용 키트를 제공한다. 상기 키트는 상기 근육전구세포의 근육세포로의 분화 정도 확인용 조성물을 포함할 수 있다. 상기 조성물에 대해서는 전술한 바와 같다. 상기 키트는 역전사 중합효소반응 키트, DNA 칩 키트, 또는 ELISA 키트일 수 있다. 상기 키트는 분석 방법에 따라 하나 이상의 다른 조성물, 용액, 또는 장치를 더 포함할 수 있다. 상기 역전사 중합효소반응 키트는 노고-에이 유전자에 특이적인 프라이머 쌍, 반응 완충액, 데옥시뉴클레오티드, Taq-폴리머라아제 및 역전사효소와 같은 효소, 및 멸균수를 포함할 수 있다. 상기 DNA 칩 키트는 노고-에이 유전자 또는 그의 단편에 해당하는 cDNA 또는 올리고뉴클레오티드가 부착되어 있는 기판 및 표지된 프로브를 제작하기 위한 시약, 제제, 및 효소를 포함할 수 있다. 상기 ELISA 키트는 노고-에이 단백질에 특이적으로 결합하는 항체를 포함할 수 있다.
다른 양상은 근육전구세포에서 근육세포로 분화 중인 세포에서 노고-에이의 발현 수준을 측정하는 단계를 포함하는, 근육전구세포의 근육세포로의 분화 정도를 확인하는 방법을 제공한다. 상기 근육전구세포 및 근육세포에 대해서는 전술한 바와 같다.
상기 발현 수준의 측정은, 노고-에이 유전자에 특이적으로 결합하는 프라이머 쌍, 프로브, 또는 노고-에이 단백질에 특이적으로 결합하는 항체를 이용하여 수행될 수 있다. 상기 프라이머, 프로브, 항체에 대해서는 전술된 바와 같다. 상기 발현 수준의 측정은 역전사효소 중합효소반응, 실시간 역전사효소 중합효소반응, 노던 블롯팅, 웨스턴 블롯팅, ELISA, 면역침전분석법, 또는 면역조직화학분석법을 이용하여 수행될 수 있다.
상기 측정 결과, 상기 분화 중인 세포에서의 노고-에이의 발현 수준이 대조군에서의 노고-에이의 발현 수준보다 높은 경우, 상기 세포는 대조군에 비해 분화가 더 진행된 세포인 것으로 결정할 수 있다. 또한, 상기 분화 중인 세포에서의 노고-에이의 발현 수준이 대조군에서의 노고-에이의 발현 수준보다 낮은 경우 상기 세포는 대조군에 비해 분화가 덜 진행된 세포인 것으로 결정할 수 있다.
다른 양상은 노고-에이 유전자의 mRNA 또는 단백질의 발현 수준을 측정하는 제제를 포함하는, 손상된 근육의 재생 정도 확인용 조성물을 제공한다. 상기 노고-에이 유전자 및 상기 제제에 대해서는 전술된 바와 같다.
다른 양상은 손상된 근육의 재생 정도 확인용 키트를 제공한다. 상기 키트는 상기 손상된 근육의 재생 정도 확인용 조성물을 포함할 수 있다. 상기 조성물에 대해서는 전술한 바와 같다. 상기 키트는 역전사 중합효소반응 키트, DNA 칩 키트, 또는 ELISA 키트일 수 있다. 각 키트에 대해서는 전술한 바와 같다.
다른 양상은 손상된 근육으로부터 분리된 세포 또는 조직에서 노고-에이의 발현 수준을 측정하는 단계를 포함하는, 손상된 근육의 재생 정도를 확인하는 방법을 제공한다.
상기 발현 수준의 측정은 노고-에이 유전자에 특이적으로 결합하는 프라이머 쌍, 프로브, 또는 노고-에이 단백질에 특이적으로 결합하는 항체를 이용하여 수행될 수 있다. 상기 프라이머, 프로브, 항체에 대해서는 전술된 바와 같다.
상기 측정 결과, 상기 손상된 근육으로부터 분리된 세포 또는 조직에서 노고-에이의 발현 수준이 대조군의 노고-에이의 발현 수준보다 낮은 경우, 상기 손상된 근육은 대조군에 비해 재생이 더 진행된 것으로 결정할 수 있다. 여기서 대조군은 손상 직후의 근육에서 분리된 세포 또는 조직일 수 있다.
다른 양상은 노고-에이 유전자의 mRNA 또는 단백질의 발현 수준을 측정하는 제제를 포함하는, 근육 손상 질환 진단용 조성물을 제공한다.
용어 "근육 손상 질환"은 근육조직 또는 근세포가 손상을 받으므로써 근육조직이 소실(loss)되는 모든 질환을 의미한다. 상기 근육 손상 질환은 근이영양증(muscular dystrophy), 근위축증(muscular atrophy), 근육염(myositis), 다발성 근육염(polymyositis), 말초혈관 질환(peripheral vascular disease), 또는 섬유증(fibrosis)일 수 있다. 상기 근이영양증은 예를 들면, 베커형 근이영양증(Becker's muscular dystrophy), 선천성 근이영양증(congenital muscular dystrophy), 뒤셴형근이영양증(Duchenne muscular dystrophy), 원위 근이영양증(distal muscular dystrophy), 에머리-드라이푸스 근이영양증(Emery-Dreifuss muscular dystrophy), 안면견갑상완 근이영양증, 지대형 근이영양증, 근긴장성 근이영양증, 안인두성 근이영양증, 근긴장성 근이영양증, 사르코글리칸증, 척수성 근육위축, 또는 BVVL(Brown-Vialetto-Van Laere syndrome)일 수 있다. 상기 노고-에이 유전자 및 상기 제제에 대해서는 전술된 바와 같다.
다른 양상은 근육 손상 질환 진단용 키트를 제공한다. 상기 키트는 상기 근육 손상 질환 진단용 조성물을 포함할 수 있다. 상기 조성물에 대해서는 전술한 바와 같다. 상기 키트는 역전사 중합효소반응 키트, DNA 칩 키트, ELISA 키트, 또는 단백질 칩 키트일 수 있다. 각 키트에 대해서는 전술한 바와 같다.
다른 양상은 개체로부터 분리된 시료에서 노고-에이 유전자의 mRNA 또는 단백질의 발현 수준을 측정하는 단계; 및 측정된 발현 수준을 대조군 시료에서의 노고-에이 유전자의 mRNA 또는 단백질의 발현 수준과 비교하는 단계를 포함하는, 개체의 근육 손상 질환을 진단하기 위한 정보제공방법을 제공한다.
상기 시료는 개체의 근육 조직으로부터 분리된 것일 수 있다. 상기 개체는 포유동물일 수 있다. 상기 포유동물은 인간, 마우스, 래트, 기니어 피그, 토끼, 원숭이, 돼지, 말, 소, 양, 영양, 개, 또는 고양이일 수 있다.
상기 근육 손상 질환, 상기 노고-에이 유전자, 상기 발현 수준의 측정에 대해서는 전술한 바와 같다. 상기 대조군 시료는 근육 손상 질환을 갖지 않는 개체일 수 있다. 개체로부터 분리된 시료에서 노고-에이 발현 수준이 대조군 시료에서 노고-에이 발현 수준에 비해 증가된 경우, 상기 개체는 근육 손상 질환을 갖는 것으로 진단될 수 있다.
노고-에이는 근육의 분화 또는 재생 정도, 또는 질환 상태를 확인하기 위해 이용될 수 있다. 또한, 본 발명은 근육의 생태병리학적인 이해를 넓히는 데에 기여할 수 있다.
도 1 및 2는 각각 분화 초기 단계 및 분화 후기 단계의 근육모세포에서 면역형광염색을 통해 노고-에이의 발현을 관찰한 결과이다.
도 3은 분화 과정 동안 노고-에이 및 다른 근육 분화 인자의 발현 양상을 나타내는 그래프이다.
도 4a는 재생 중인 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다.
도 4b는 질환 상태의 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다.
도 5는 재생 중인 근육 및 질환 상태의 근육에서 노고-에이의 발현 수준을 q-PCR을 이용하여 분석한 결과를 나타낸다.
도 3은 분화 과정 동안 노고-에이 및 다른 근육 분화 인자의 발현 양상을 나타내는 그래프이다.
도 4a는 재생 중인 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다.
도 4b는 질환 상태의 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다.
도 5는 재생 중인 근육 및 질환 상태의 근육에서 노고-에이의 발현 수준을 q-PCR을 이용하여 분석한 결과를 나타낸다.
이하, 실시예를 통하여 본 발명을 더욱 상세히 설명하고자 한다. 이들 실시예는 오로지 본 발명을 보다 구체적으로 설명하기 위한 것으로, 본 발명의 범위가 이들 실시예에 의해 제한되는 것은 아니다.
실시예
1:
근육모세포의
분화 과정 중 노고-에이 발현 분석
1.
근육모세포의
분화 유도
근육세포의 분화 실험을 하기 위해 마우스 유래 근육모세포주 C2C12를 사용하였다 (ATCC®, 미국). 고농도 글루코스-Dulbecco's Modified Eagle Medium (H-DMEM, Life technologies, 미국), 10% 우태혈청 (FBS, ATLAS, 미국), 및 1% 페니실린-스트렙토마이신 (PS, Life technologies, 미국)이 함유된 일반증식조건 배지에서 증식을 유지하였다. 그 후 세포수가 80% 이상이 되었을 때, 저농도 글루코스-DMEM (L-DMEM, GE healthcare, 미국), 2% 말 혈청 (HS, Life technologies, 미국), 및 1% PS (Life technologies, 미국)이 함유된 분화조건 배지로 교체하여 분화를 유도하였다. 37℃, 5% CO2 조건으로 인큐베이터에서 배양하였다. 하기와 같이 근육모세포의 분화 과정 동안의 노고-에이의 발현을 분석하였다.
2.
면역형광염색을
이용한 노고-에이 발현 분석
분화 조건 배지로의 교체 전과 교체 14일 후, 즉, 분화 초기 단계와 후기 단계의 근육모세포에 대하여 면역형광염색을 실시하였다. 근육 분화 단계를 대조하기 위해 Pax7, MyoD, Myogenin, MYH2 단백질을 같이 동시-염색하였다.
배양된 세포를 4% 파라포름알데히드 (Sigma, 미국)로 15분간 실온에서 고정한 후 0.01M 인산완충식염수 (PBS, Gendepot, 미국)로 5분간 3회 수세하였다. 그 후 고정된 세포를 메탄올로 -20℃에서 10분간 인큐베이션하였다. PBS로 5분간 3회 수세 후, 0.01M PBS 중 3% 소혈청알부민 (Sigma, 미국)으로 실온에서 1시간 블로킹하였다. 노고-에이 항체 (1:500; Abcam, 미국), Pax7 (1:50; Developmental Studies Hybridoma Bank, 미국), MyoD (1:200; Santacruz, 미국), Myogenin (1:20; DSHB, 미국), 및 MYH2 (1:200; Santacruz, 미국)를 각각 0.1% Triton X-100 (Sigma, 미국)이 첨가된 PBS (PBS/T)에 희석하였다. 준비된 세포들을 각각의 항체 희석액으로 4℃에 약 18시간 동안 염색하였다. PBS로 5분 3회 수세 후, 플루오레신 이소시아네이트 (FITC, 1:500; Life technologies, 미국) 및 테트라메틸로다민 (TRITC, 1:500; Life technologies, 미국)이 첨가된 PBS/T로 1시간 동안 실온에서 염색하였다. PBS로 5분간 3회 수세 후, 4',6-디아미디노-2-페닐인돌 (DAPI, 0.1 ㎍/ml; Sigma, 미국)로 최종 염색하였다. 공초점 레이저 주사 현미경 (Carl zeiss, 독일)을 이용하여 사진 촬영을 하였다. Zen 2009 (Carl zeiss,독일) 소프트웨어로 이미지를 분석하였다.
도 1 및 2는 각각 분화 초기 단계 및 분화 후기 단계의 근육모세포에서 면역형광염색을 통해 노고-에이의 발현을 관찰한 결과이다. 도 1 및 2에 나타낸 바와 같이, 노고-에이의 발현은 분화된 근육세포인 근육대롱(myotube)의 표지 단백질인 MYH2의 발현과 상당히 일치하였다. 이에 비해, 노고-에이의 발현은 분화 중인 근육세포 표지 단백질인 MyoD와 Myogenin과는 부분적으로 일치하고, 근육모세포 표지 단백질인 PAX7과는 일치하지 않았다.
3.
RT
-
PCR
및 q-
PCR
을 이용한 노고-에이 발현 분석
분화 유도된 근육모세포로부터 RNA를 1일 간격으로 추출하여 RT-PCR 혹은 q-PCR로 노고-에이의 발현을 관찰하였다
PCR 반응 조건은 다음과 같았다. AccuPower® PCR premix (Bioneer, 대한민국)에 조건당 노고-에이 프라이머 세트 (서열번호 3 및 4) 및 Gapdh 프라이머 세트 (서열번호 5 및 6) 각 1 ㎕, 1 ㎕ cDNA, 및 17 ㎕ 초순수를 혼합하였다. Biorad 사의 T100 기기를 이용하여, 95℃에서 4분 동안 반응 후, 95℃에서 30초, 58℃에서 30초, 및 72℃에서 30초로 35회 반복하고, 72℃에서 7분 동안 반응시켰다.
qPCR 반응 조건은 다음과 같았다. 2X SYBR green master mix (TAKARA, 미국) 12.5 ㎕, 프라이머 세트 각 2.5 ㎕, cDNA 3 ㎕, 및 DEPC water 7 ㎕을 25 ㎕ 반응 용량으로 혼합하였다. 프라이머 세트는 노고-에이 (서열번호 7 및 8), Myogenin (Qiagen, 미국), MyoD (Qiagen, 미국), Gapdh (Qiagen, 미국)를 이용하였다. Qiagen 사의 Rotor-Gene Q를 이용하여 95℃에서 5분 동안 반응 후, 95℃에서 30초, 및 60℃에서 10초로 40회 반복하여 반응시켰다.
도 3은 분화 과정 동안 노고-에이 및 다른 근육 분화 인자의 발현 양상을 나타내는 그래프이다. 도 3에 나타낸 바와 같이, 분화가 진행됨에 따라 노고-에이 유전자 또한 Myogenin과 MyoD와 더불어 그 발현이 지속적으로 증가하는 경향을 나타내었다.
실시예
2: 근육의 재생 과정에서 노고-
에이의
발현 분석
1. 근육의 재생 과정 유도
근육의 재생 과정을 유도하기 위하여, Antonio S.J. Lee, BioArchithecture, 3:2, 25-37; 2013에 기재된 방법에 따라, 미오톡신(myotoxin)인 노텍신(notexin)을 근육에 주입하여 근육 손상을 유도하였다. 8주령 된 수컷 C57BL/6 마우스(n=9) (중앙실험동물, 대한민국)의 양쪽 하지 피부를 면도한 후, 정강이뼈 옆에 있는 앞정강근에 주사기로 노텍신 (Sigma-aldrich, 미국)을 5 mg/l로 생리식염수에 희석하여 한 다리당 20 ㎕씩 주입하였다 (0.1 ㎍/1 leg).
2.
RT
-
PCR
및 q-
PCR
을 이용한 노고-에이 발현 분석
노텍신 주입 후 6일, 9일, 및 12일차에 각각 3마리, 3마리, 및 2마리씩 부검하여 근육을 분리하였다. 분리된 근육 시료로부터 RNA를 추출하여 RT-PCR 혹은 q-PCR로 노고-에이의 발현을 관찰하였다.
도 4a는 재생 중인 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다. 음성 대조군인 온전한 근육 및 양성 대조군인 뇌조직은 각각 손상이 유도되지 않은 8주령 된 수컷 C57BL/6 마우스에서 분리된 근육 시료 및 뇌 조직이다.
도 5는 재생 중인 근육에서 노고-에이의 발현 수준을 q-PCR을 이용하여 분석한 결과를 나타낸다. 도 5에 나타낸 바와 같이, 손상된 근육의 재생이 진행됨에 따라 노고-에이의 발현이 감소하고, 손상 받지 않은 근육의 발현량에 가까워지는 경향을 나타내었다.
실시예
3: 질환 상태의 근육에서 노고-
에이의
발현 분석
질환 동물 모델인 3개월령, 6개월령, 또는 15개월령 수컷 MDX 마우스(n=3)에서(Jacques P.Tremblay Lab에서 분양), 실시예 2와 동일한 방법으로 근육 내 노고-에이의 발현을 RT-PCR 과 q-PCR을 이용하여 분석하였다. 대조군으로는 1개월령, 3개월령, 6개월령, 또는 15개월령된 수컷 C57BL/6 마우스와 그의 뇌 조직을 이용하였다.
도 4b는 질환 상태의 근육에서 노고-에이의 발현을 RT-PCR을 이용하여 분석한 결과이다. 도 4b에 나타낸 바와 같이, 질환 상태의 근육에서 노고-에이의 발현 수준은 질환을 갖지 않는 대조군 근육에 비하여 높게 나타났다.
도 5는 질환 상태의 근육에서 노고-에이의 발현 수준을 q-PCR을 이용하여 분석한 결과를 나타낸다. 도 5에 나타낸 바와 같이, 질환을 갖는 근육에서 노고-에이의 발현은 질환을 갖지 않는 대조군의 근육에 비하여 높게 나타나는 것으로 확인되었다.
<110> Kyungpook National University industry academic cooperation foundation
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disease condition of muscle
<130> PN107894
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<170> KopatentIn 2.0
<210> 1
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Met Glu Asp Ile Asp Gln Ser Ser Leu Val Ser Ser Ser Ala Asp Ser
1 5 10 15
Pro Pro Arg Pro Pro Pro Ala Phe Lys Tyr Gln Phe Val Thr Glu Pro
20 25 30
Glu Asp Glu Glu Asp Glu Glu Asp Glu Glu Glu Glu Glu Asp Asp Glu
35 40 45
Asp Leu Glu Glu Leu Glu Val Leu Glu Arg Lys Pro Ala Ala Gly Leu
50 55 60
Ser Ala Ala Pro Val Pro Pro Ala Ala Ala Pro Leu Leu Asp Phe Ser
65 70 75 80
Ser Asp Ser Val Pro Pro Ala Pro Arg Gly Pro Leu Pro Ala Ala Pro
85 90 95
Pro Thr Ala Pro Glu Arg Gln Pro Ser Trp Glu Arg Ser Pro Ala Ala
100 105 110
Ser Ala Pro Ser Leu Pro Pro Ala Ala Ala Val Leu Pro Ser Lys Leu
115 120 125
Pro Glu Asp Asp Glu Pro Pro Ala Arg Pro Pro Ala Pro Ala Gly Ala
130 135 140
Ser Pro Leu Ala Glu Pro Ala Ala Pro Pro Ser Thr Pro Ala Ala Pro
145 150 155 160
Lys Arg Arg Gly Ser Gly Ser Val Asp Glu Thr Leu Phe Ala Leu Pro
165 170 175
Ala Ala Ser Glu Pro Val Ile Pro Ser Ser Ala Glu Lys Ile Met Asp
180 185 190
Leu Lys Glu Gln Pro Gly Asn Thr Val Ser Ser Gly Gln Glu Asp Phe
195 200 205
Pro Ser Val Leu Phe Glu Thr Ala Ala Ser Leu Pro Ser Leu Ser Pro
210 215 220
Leu Ser Thr Val Ser Phe Lys Glu His Gly Tyr Leu Gly Asn Leu Ser
225 230 235 240
Ala Val Ala Ser Thr Glu Gly Thr Ile Glu Glu Thr Leu Asn Glu Ala
245 250 255
Ser Arg Glu Leu Pro Glu Arg Ala Thr Asn Pro Phe Val Asn Arg Glu
260 265 270
Ser Ala Glu Phe Ser Val Leu Glu Tyr Ser Glu Met Gly Ser Ser Phe
275 280 285
Asn Gly Ser Pro Lys Gly Glu Ser Ala Met Leu Val Glu Asn Thr Lys
290 295 300
Glu Glu Val Ile Val Arg Ser Lys Asp Lys Glu Asp Leu Val Cys Ser
305 310 315 320
Ala Ala Leu His Asn Pro Gln Glu Ser Pro Ala Thr Leu Thr Lys Val
325 330 335
Val Lys Glu Asp Gly Val Met Ser Pro Glu Lys Thr Met Asp Ile Phe
340 345 350
Asn Glu Met Lys Met Ser Val Val Ala Pro Val Arg Glu Glu Tyr Ala
355 360 365
Asp Phe Lys Pro Phe Glu Gln Ala Trp Glu Val Lys Asp Thr Tyr Glu
370 375 380
Gly Ser Arg Asp Val Leu Ala Ala Arg Ala Asn Met Glu Ser Lys Val
385 390 395 400
Asp Lys Lys Cys Phe Glu Asp Ser Leu Glu Gln Lys Gly His Gly Lys
405 410 415
Asp Ser Glu Ser Arg Asn Glu Asn Ala Ser Phe Pro Arg Thr Pro Glu
420 425 430
Leu Val Lys Asp Gly Ser Arg Ala Tyr Ile Thr Cys Asp Ser Phe Ser
435 440 445
Ser Ala Thr Glu Ser Thr Ala Ala Asn Ile Phe Pro Val Leu Glu Asp
450 455 460
His Thr Ser Glu Asn Lys Thr Asp Glu Lys Lys Ile Glu Glu Arg Lys
465 470 475 480
Ala Gln Ile Ile Thr Glu Lys Thr Ser Pro Lys Thr Ser Asn Pro Phe
485 490 495
Leu Val Ala Ile His Asp Ser Glu Ala Asp Tyr Val Thr Thr Asp Asn
500 505 510
Leu Ser Lys Val Thr Glu Ala Val Val Ala Thr Met Pro Glu Gly Leu
515 520 525
Thr Pro Asp Leu Val Gln Glu Ala Cys Glu Ser Glu Leu Asn Glu Ala
530 535 540
Thr Gly Thr Lys Ile Ala Tyr Glu Thr Lys Val Asp Leu Val Gln Thr
545 550 555 560
Ser Glu Ala Ile Gln Glu Ser Ile Tyr Pro Thr Ala Gln Leu Cys Pro
565 570 575
Ser Phe Glu Glu Ala Glu Ala Thr Pro Ser Pro Val Leu Pro Asp Ile
580 585 590
Val Met Glu Ala Pro Leu Asn Ser Leu Leu Pro Ser Thr Gly Ala Ser
595 600 605
Val Ala Gln Pro Ser Ala Ser Pro Leu Glu Val Pro Ser Pro Val Ser
610 615 620
Tyr Asp Gly Ile Lys Leu Glu Pro Glu Asn Pro Pro Pro Tyr Glu Glu
625 630 635 640
Ala Met Ser Val Ala Leu Lys Thr Ser Asp Ser Lys Glu Glu Ile Lys
645 650 655
Glu Pro Glu Ser Phe Asn Ala Ala Ala Gln Glu Ala Glu Ala Pro Tyr
660 665 670
Ile Ser Ile Ala Cys Asp Leu Ile Lys Glu Thr Lys Leu Ser Thr Glu
675 680 685
Pro Ser Pro Glu Phe Ser Asn Tyr Ser Glu Ile Ala Lys Phe Glu Lys
690 695 700
Ser Val Pro Asp His Cys Glu Leu Val Asp Asp Ser Ser Pro Glu Ser
705 710 715 720
Glu Pro Val Asp Leu Phe Ser Asp Asp Ser Ile Pro Glu Val Pro Gln
725 730 735
Thr Gln Glu Glu Ala Val Met Leu Met Lys Glu Ser Leu Thr Glu Val
740 745 750
Ser Glu Thr Val Thr Gln His Lys His Lys Glu Arg Leu Ser Ala Ser
755 760 765
Pro Gln Glu Val Gly Lys Pro Tyr Leu Glu Ser Phe Gln Pro Asn Leu
770 775 780
His Ile Thr Lys Asp Ala Ala Ser Asn Glu Ile Pro Thr Leu Thr Lys
785 790 795 800
Lys Glu Thr Ile Ser Leu Gln Met Glu Glu Phe Asn Thr Ala Ile Tyr
805 810 815
Ser Asn Asp Asp Leu Leu Ser Ser Lys Glu Asp Lys Met Lys Glu Ser
820 825 830
Glu Thr Phe Ser Asp Ser Ser Pro Ile Glu Ile Ile Asp Glu Phe Pro
835 840 845
Thr Phe Val Ser Ala Lys Asp Asp Ser Pro Lys Glu Tyr Thr Asp Leu
850 855 860
Glu Val Ser Asn Lys Ser Glu Ile Ala Asn Val Gln Ser Gly Ala Asn
865 870 875 880
Ser Leu Pro Cys Ser Glu Leu Pro Cys Asp Leu Ser Phe Lys Asn Thr
885 890 895
Tyr Pro Lys Asp Glu Ala His Val Ser Asp Glu Phe Ser Lys Ser Arg
900 905 910
Ser Ser Val Ser Lys Val Pro Leu Leu Leu Pro Asn Val Ser Ala Leu
915 920 925
Glu Ser Gln Ile Glu Met Gly Asn Ile Val Lys Pro Lys Val Leu Thr
930 935 940
Lys Glu Ala Glu Glu Lys Leu Pro Ser Asp Thr Glu Lys Glu Asp Arg
945 950 955 960
Ser Leu Thr Ala Val Leu Ser Ala Glu Leu Asn Lys Thr Ser Val Val
965 970 975
Asp Leu Leu Tyr Trp Arg Asp Ile Lys Lys Thr Gly Val Val Phe Gly
980 985 990
Ala Ser Leu Phe Leu Leu Leu Ser Leu Thr Val Phe Ser Ile Val Ser
995 1000 1005
Val Thr Ala Tyr Ile Ala Leu Ala Leu Leu Ser Val Thr Ile Ser Phe
1010 1015 1020
Arg Ile Tyr Lys Gly Val Ile Gln Ala Ile Gln Lys Ser Asp Glu Gly
1025 1030 1035 1040
His Pro Phe Arg Ala Tyr Leu Glu Ser Glu Val Ala Ile Ser Glu Glu
1045 1050 1055
Leu Val Gln Lys Tyr Ser Asn Ser Ala Leu Gly His Val Asn Ser Thr
1060 1065 1070
Ile Lys Glu Leu Arg Arg Leu Phe Leu Val Asp Asp Leu Val Asp Ser
1075 1080 1085
Leu Lys Phe Ala Val Leu Met Trp Val Phe Thr Tyr Val Gly Ala Leu
1090 1095 1100
Phe Asn Gly Leu Thr Leu Leu Ile Leu Ala Leu Ile Ser Leu Phe Ser
1105 1110 1115 1120
Ile Pro Val Ile Tyr Glu Arg His Gln Ala Gln Ile Asp His Tyr Leu
1125 1130 1135
Gly Leu Ala Asn Lys Ser Val Lys Asp Ala Met Ala Lys Ile Gln Ala
1140 1145 1150
Lys Ile Pro Gly Leu Lys Arg Lys Ala Glu
1155 1160
<210> 2
<211> 6165
<212> RNA
<213> Mus musculus
<400> 2
cccctgccgg gaaggtgagt cacgccaaac tgggcggaga gtctgccctc ctctctcagt 60
tgctcatctg ggcggcggcg gctgctgcaa ctgaggacag ggcgggtggc gcatctcgag 120
cgcggaggca ggaggagaag tcttattgtt cctggagctg tcgcctttgc gggttcctcg 180
gcttggttcg gccagcccgg cctctgccag tcctgcccaa cccccacaac cgcccgcggc 240
tctgaggaga agtggcccgc ggcggcagta gctgcagcat catcgccgac catggaagac 300
atagaccagt cgtcgctggt ctcctcgtcc gcggatagcc cgccccggcc cccgcccgct 360
ttcaagtacc agttcgtgac ggagcccgag gacgaggagg acgaggaaga cgaggaggag 420
gaggaggacg acgaggacct ggaggaattg gaggtgctgg agaggaagcc cgcagccggg 480
ctgtccgcgg ctccggtgcc ccccgccgcc gcaccgctgc tggacttcag cagcgactcg 540
gtgccccccg cgccccgcgg gccgctgccg gccgcgcccc ccaccgcccc tgagaggcag 600
ccgtcctggg aacgcagccc cgcggcgtcc gcgccatccc tgccgcccgc tgccgcagtc 660
ctgccctcca agctcccgga ggacgacgag cctccagcgc ggcctccggc gccagccggc 720
gcgagccccc tagcggagcc cgccgcgccc ccttccacgc cggccgcgcc caagcgcagg 780
ggctcgggct cagtggatga gacccttttt gctcttcctg ctgcatctga gcctgtgata 840
ccctcctctg cagaaaaaat tatggatttg aaggagcagc caggtaacac tgtttcgtct 900
ggtcaagagg atttcccatc tgtcctgttt gaaactgctg cctctcttcc ttctctatct 960
cctctctcaa ctgtttcttt taaagaacac ggataccttg gtaacttatc agcagtggca 1020
tccacagaag gaactattga agaaacttta aatgaagctt ctagagaatt gccagagagg 1080
gcaacaaatc catttgtaaa tagagagtca gcagagtttt cagtattaga atactcagaa 1140
atgggatcat ctttcaatgg ctccccaaaa ggagagtcag ccatgttagt agaaaacact 1200
aaggaagaag taattgtgag gagtaaagac aaagaggatt tagtttgtag tgcagccctt 1260
cataatccac aagagtcacc tgcgaccctt actaaagtgg ttaaagaaga cggagttatg 1320
tctccagaaa agacaatgga catttttaat gaaatgaaaa tgtcagtggt agcacctgtg 1380
agggaagagt atgcagattt taagccattt gaacaagcat gggaagtgaa agatacttat 1440
gagggaagta gggatgtgct ggctgctaga gctaatatgg aaagtaaagt ggacaaaaaa 1500
tgctttgaag atagcctgga gcaaaaaggt catgggaagg atagtgaaag cagaaatgag 1560
aatgcttctt tccccaggac cccagaactt gtgaaggacg gctccagagc gtacatcacc 1620
tgtgattcct ttagctcagc aaccgagagt actgcagcaa acattttccc tgtgctagaa 1680
gatcacactt cagaaaacaa aacagatgaa aaaaaaatag aagaaaggaa ggcccaaatt 1740
ataacagaga agactagccc caaaacgtca aatcctttcc ttgtagcaat acatgattct 1800
gaggcagatt atgtcacaac agataattta tcaaaggtga ctgaggcagt agtggcaacc 1860
atgcctgaag gtctaacgcc agatttagtt caggaagcat gtgaaagtga actgaacgaa 1920
gccacaggta caaagattgc ttatgaaaca aaagtggact tggtccagac atcagaagct 1980
atacaagagt caatttaccc cacagcacag ctttgcccat catttgagga agctgaagca 2040
actccgtcac cagttttgcc tgatattgtt atggaagcgc cattaaattc tctccttcca 2100
agcactggtg cttctgtagc gcagcccagt gcatccccac tagaagtacc gtctccagtt 2160
agttatgacg gtataaagct tgagcctgaa aatcccccac catatgaaga agccatgagt 2220
gtagcactaa aaacatcgga ctcaaaggaa gaaattaaag agcctgaaag ttttaatgca 2280
gctgctcagg aagcagaagc tccttatata tccattgcat gtgatttaat taaagaaaca 2340
aagctctcca ctgagccaag tccagagttc tctaattatt cagaaatagc aaaatttgag 2400
aagtcggtgc ctgatcactg tgagctcgtg gatgattcct cacccgaatc tgaaccagtt 2460
gacttattta gtgatgattc aattcctgaa gtcccacaaa cacaagagga ggctgtgatg 2520
ctaatgaagg agagtctcac tgaagtgtct gagacagtaa cacaacacaa acataaggag 2580
agacttagtg cttcacctca ggaggtagga aagccatatt tagagtcttt tcagcccaat 2640
ttacatatta caaaagatgc tgcatctaat gaaattccaa cattgaccaa aaaggagaca 2700
atttctttgc aaatggaaga gtttaatact gcaatttatt ccaatgatga cttactttct 2760
tctaaggaag acaaaatgaa agaaagtgaa acattttccg attcatctcc cattgagata 2820
atagatgagt ttcccacatt tgtcagtgct aaagatgatt ctcctaagga gtacactgac 2880
ctagaagtat ccaacaaaag tgaaattgct aatgtccaga gcggggccaa ttcgttgcct 2940
tgctcagaat tgccctgtga cctttctttc aagaatacat atcctaaaga tgaagcacat 3000
gtctcagatg aattctccaa aagtaggtcc agtgtatcta aggtgccctt attgcttcca 3060
aatgtttctg ctttggaatc tcaaatagaa atgggcaaca tagttaaacc caaagtactt 3120
acgaaagaag cagaggaaaa acttccttct gatacagaga aagaggacag atccctgaca 3180
gctgtattgt cagcagagct gaataaaact tcagttgttg acctcctgta ctggagagac 3240
attaagaaga ctggagtggt gtttggtgcc agcttattcc tgctgctgtc tctgacagtg 3300
ttcagcattg tcagtgtaac ggcctacatt gccttggccc tgctctctgt gactatcagc 3360
tttaggatat ataagggtgt gatccaagct atccagaaat cagatgaagg ccacccattc 3420
agggcatatt tggaatctga agttgccata tcagaggaat tggttcagaa atatagtaat 3480
tctgctcttg gtcatgtgaa cagcacaata aaagaattga ggcgtctctt cttagttgat 3540
gatttagttg attccctgaa gtttgcagtg ttgatgtggg tatttactta cgttggtgcc 3600
ttgttcaatg gtttgacact actgatttta gctctgatct cactcttcag tattcctgtt 3660
atatatgaac ggcatcaggc gcagatagat cattatctag gacttgcaaa caagagcgtt 3720
aaggatgcca tggccaaaat ccaagcaaaa atccctggat tgaagcgcaa agcagaatga 3780
aaaggcccca aacagtagac attcatcttt aaaggggaca ctcccttggt tacggggtgg 3840
gcgggtcagg ggtgagccct gggtggccgt gcagtttcag ttatttttag cagtgcactg 3900
tttgaggaaa aattacctgt cttgacttcc tgtgtttatc atcttaagta ttgtaagctg 3960
ctgtgtatgg atctcattgt agtcatactt gttttcccca gatgaggcac ttggtgaata 4020
aaggatgctg ggaaaactgt gtgttatatt ctgttgcagg tagtctggct gtatttggaa 4080
agttgcaaag aaggtagatt tgggggcagg aaaaacaacc cttttcacag tgtactgtgt 4140
ttggttggtg taaaactgat gcagattttt ctgaaatgag atgtttagat gagcatacta 4200
ctaaagcaga gtggaaaaat ctgtctttat ggtatgttct aggtgtattg tgatttactg 4260
ttagattgcc aatataagta aatatagaca taatctatat atagtgtttc acaaagctta 4320
gatctttaac cttgcagctg ccccacagtg cttgacctct gagtcattgg ttatacagtg 4380
tagtcccaag cacataaact aggaagagaa tgtatttgta ggagcgctac cacctgtttt 4440
caagagaaca tagaactcca acgtaaccgt catttcaaag atttactgta tgtatagttg 4500
attttgtgga ctgaatttaa tgcttccaaa tgtttgcagt taccaaacat tgttatgcaa 4560
gaaatcataa aatgaagact tataccattg tgtttaagct gtattgaatt atctgtggaa 4620
tgcattgtga actgtaaagc aaagtatcaa taaagcttat agacttacct ttgtgtttag 4680
tgttttagtt tcatgaatgc acagcaaaaa cacggtggta ggcttagaga gtggacacat 4740
ggtaacatgc ttttagaaag gttttagttc atgaaacagc ttaagaacaa agaatatatt 4800
tacatagtga gatttatttg actcataaca aaaggtttta aattatttta tactttgaaa 4860
ataaattcat gcaccaatat tttaacagaa tacagtgcaa gatttatgaa tatacataaa 4920
attacaccat ataaatttta caaataagac tttcaaagtc tttataacag acactattgc 4980
tcttcaaata tatacatata tcattgatta gtcagttgtt catccacatg gttacttaat 5040
gcaagatctg tctgaatgaa atgtcagtag tacaagacag gcagacacag tgatcactca 5100
gcatcaccag gtacagaaaa cagaatcagg gctgcatagg gctctactga ggacccagca 5160
acctgctagt gggttgatgt aaggcattaa taagttggcg tgtaaaatag cttaatgtgt 5220
aatctaattc ttttagaatg ctgaagcact tctgtggtaa atgttgataa tctattcttt 5280
aactgaaaat gcttatttca acctttctct aaaatggcaa cttcatataa ctagaaactc 5340
aaggtctaga attttagtgc acagactgga aggactcggt ggtgtgtgta ctcacgactc 5400
caactcccat cagccttctt aactaatagt cgtcaagtca cattctgtcc gacaactgac 5460
tggagaaact caaatactcc ttacagtggg gaatatgttc aagaggtttt taaaaatctg 5520
aatttaccct gcattaatca tctgaaatga gcagagccaa gccagtccta cccaagaggg 5580
tgtaaactaa acagcaagac agctcactcg tcacactgga gctttccctg ctttccgtgt 5640
tgctactttc tgtggagctg gactcttctt tgctgctcac cttataactg ctttcctaga 5700
gcaggacata gtggtgaatt tgctaatccc atagccctcc tcagttttga agttttagca 5760
ccatgtgaag gaagaagacg acatgggagg tgaggcagca gcccacacaa ctgggaactt 5820
tgaaggcact taatggatat aaaactgcaa atgaaacttt taaaaattaa cattttaaaa 5880
cttgtttata catggctcat ttagttttga aagctaaatg tggcaaacag ggtatttctg 5940
tacttaagtg cttccaactt acattgtgtc cagtgaacat tcttaaaata catagaaaca 6000
gaatagcaaa acacctttgt aaaagtctta atgcaaatta aacgctacat attggttagg 6060
gtaattgagg atgtaaattt tatctgtggg ctacatcatc ccaattttgc cgttagtgaa 6120
gttacaataa gtataggttt tagtctccag aacagaagca aacag 6165
<210> 3
<211> 19
<212> DNA
<213> Artificial Sequence
<220>
<223> NogoA F
<400> 3
caagctcccg gaggacgac 19
<210> 4
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> NogoA R
<400> 4
gccctctctg gcaattctct 20
<210> 5
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Gapdh F
<400> 5
actcacggaa aattcaacgg 20
<210> 6
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> Gapdh R
<400> 6
accagtggat gcagggatga 20
<210> 7
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<223> qNogoA F
<400> 7
ctcagtggat gagacccttt ttgc 24
<210> 8
<211> 21
<212> DNA
<213> Artificial Sequence
<220>
<223> qNogoA R
<400> 8
cagtgttacc tggctgctcc t 21
Claims (20)
- 노고-에이(Nogo-A) 유전자의 mRNA 또는 단백질의 발현 수준을 측정하는 제제를 포함하는, 근육전구세포의 근육세포로의 분화 정도 확인용 조성물.
- 청구항 1에 있어서, 상기 유전자는 서열번호 1의 폴리펩티드를 코딩하는 것인 조성물.
- 청구항 1에 있어서, 상기 제제는 상기 유전자에 특이적으로 결합하는 프라이머 쌍, 프로브, 또는 상기 단백질에 특이적으로 결합하는 항체인 것인 조성물.
- 청구항 1 내지 3 중 어느 한 항의 조성물을 포함하는, 근육전구세포의 근육세포로의 분화 정도 확인용 키트.
- 청구항 4에 있어서, 상기 키트는 역전사 중합효소반응 키트, DNA 칩 키트, ELISA 키트, 또는 단백질 칩 키트인 것인 키트.
- 근육전구세포에서 근육세포로 분화 중인 세포에서 노고-에이의 발현 수준을 측정하는 단계를 포함하는, 근육전구세포의 근육세포로의 분화 정도를 확인하는 방법.
- 청구항 6에 있어서, 상기 발현 수준의 측정은 노고-에이 유전자에 특이적으로 결합하는 프라이머 쌍, 프로브, 또는 노고-에이 단백질에 특이적으로 결합하는 항체를 이용하는 것인 방법.
- 청구항 6에 있어서, 상기 분화 중인 세포의 노고-에이의 발현 수준이, 대조군의 노고-에이의 발현 수준보다 높은 경우 상기 분화 중인 세포는 대조군에 비해 분화가 더 진행된 세포인 것으로 결정하고, 대조군의 노고-에이의 발현 수준보다 낮은 경우 상기 분화 중인 세포는 대조군에 비해 분화가 덜 진행된 세포인 것으로 결정하는 것인 방법.
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
- 삭제
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| KR102279751B1 (ko) * | 2019-06-28 | 2021-07-21 | 경북대학교 산학협력단 | 근원성 인자 또는 nogo-a 측정 제제를 포함하는 근육병 진단용 조성물과 이를 이용한 근육병 진단 방법 |
| KR102409771B1 (ko) | 2020-06-17 | 2022-06-16 | 경북대학교 산학협력단 | Nogo-A 및 필라민-C의 상호 결합을 이용한 근육질환의 치료제 스크리닝 방법 |
Citations (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| JP2011173899A (ja) | 2002-08-10 | 2011-09-08 | Yale Univ | Ｎｏｇｏ受容体アンタゴニスト |
| JP2012213406A (ja) | 1998-11-06 | 2012-11-08 | Univ Of Zurich | Ｎｏｇｏ遺伝子のヌクレオチド配列およびタンパク質配列、ならびにそれに基づく方法 |
| US20130316933A1 (en) * | 2010-12-09 | 2013-11-28 | Universidad De Zaragoza | Method for the diagnosis, prognosis and monitoring of muscular degeneration |
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Patent Citations (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| JP2012213406A (ja) | 1998-11-06 | 2012-11-08 | Univ Of Zurich | Ｎｏｇｏ遺伝子のヌクレオチド配列およびタンパク質配列、ならびにそれに基づく方法 |
| JP2011173899A (ja) | 2002-08-10 | 2011-09-08 | Yale Univ | Ｎｏｇｏ受容体アンタゴニスト |
| US20130316933A1 (en) * | 2010-12-09 | 2013-11-28 | Universidad De Zaragoza | Method for the diagnosis, prognosis and monitoring of muscular degeneration |
Non-Patent Citations (1)
| Title |
|---|
| The Journal of Neuroscience, Vol. 22, No. 13, pp. 5505-5515 (2002. 07.01.) |
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