KR900008742B1 - 핵산 서열의 증대 방법 - Google Patents
핵산 서열의 증대 방법 Download PDFInfo
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Abstract
내용 없음.
Description
제1도는 증대시키려고 하는 인체 β-글로빈의 94개 염기쌍 서열을 나타낸 것이고, 겸상 적혈구 빈혈증(sickle cell anemia)과 관련된 단일 염기쌍의 변화는 94-머(94-mer) 아래에 표시되어 있다.
제2도는 인체의 야생형 DNA에 및 정상적인 β-글로빈 유전자의 1.9kbH I 단편을 함유하는 플라스미드(pBR 328 : HbA로 표기)에 함유되어 있는 94-머(mer)의 증대를 제시하는 폴리아크릴 아미드 겔 전기 영동의 자동방사선 사진다.
제3도는 pBR 328-HbA 및 β-글로빈의 겸상적혈구 대립인자(allele)의 1.9kbH I 단편을 함유하는 플라스미드(pBR 328 : HbS로 표기)[durlt에서, pBR 328 : HbA에서 증대되는 서열은II로 절단되지만, pBR 328 : HbS에 증대되는 서열은 동일한 처리를 하여도II로 절단되지 않는다]에 존재하는 어떤 특정의 표적 94-머 서열의 증대를 설명해주는 폴리아크릴아미드 겔 전기영동의 자동방사선 사진이다.
제4도는 인체 β-글로빈의 목적하는 94-머 서열의 증대를 위한 폴리머라제 연쇄 반응(polymerase chain reaction)의 단계 및 생성물을 2개의 올리고뉴클레오타이드 프라이머(oligonucleotide primers)를 사용하여 3개의 사이클로 상세하게 설명한 것이다.
제5도는 pBR 328 : HbA에서 240-머 서열의 4개의 사이클 후의 증대를 설명해주는, 폴리아크릴아미드겔 전기영동의 자동방사선 사진이다.[여기에서, 분취량(aliquots)은I(레인 3),II(레인 4) 또는f I(레인 5)로 분해시킨다]. 레인 1은 분자량 표준이고 레인 2는 무상(intact) 240-bp 생성물을 함유한다.
제6도는I 및f I 제한부의 범위에서 정상적인 β-글로빈 유전자(βA) 및 경상 적혈구 세포의 β-글로빈 유전자(βs)의 서열을 보여준다. [여기에서, βA에 있어 외줄은I 부위(CTGAG)의 위치를 나타내고 βA 및 βS에 있어 겹줄은 Hinf I 부위(GACTC)의 위치를 나타낸다].
제9도는 본 발명에 의해 증대시킨 전 인체 DNA의 샘플에 베타-글로빈 대립인자가 존재함을 특징으로 하는 제7도에서와 동일한 40-머 탐침의 용도를 설명하기 위한 폴리아크릴아미드 겔 전기영동의 자동방사선 사진이다.
제10도는 에티디움 브로마이드 및 UV 광을 사용하여 가시화한 6% 누시이브 아가로오스 겔(Nusieve agarose gel)의 사진이다. 이 사진은 110-bp 증대 생성물의 아단편의 증대[여기에서, 아단편은 110-bp 단편 이내에서 내부에 존재하는 세트(inner nested set)이다]를 설명한 것이다.
본 발명은 현존하는 핵산 서열을 증대시키는 방법에 관한 것이다. 특히 본 발명은 주어진 DNA 또는 RNA의 서열로부터 특정 서열을 초기에 존재하는 양과 비해서 다량으로 생산하는 방법에 관한 것이다. DNA 또는 RNA는 일본쇄 또는 이본쇄 일 수 있으며, 비교적 순수한 종 또는 핵산 혼합물의 성분일 수 있다. 본 발명의 방법을 목적하는 핵산 서열의 증대를 달성하기 위한 반복 반응에 이용한다.
특히 진단적 적용(diagnoistic application)을 위해 표적의 핵산 서열은 해당 DNA 또는 RNA의 단지 소부분(small portion)일 수 있으며, 이 경우 비동위적으로 표지(nonisotopically labeled)하거나 말단을 표지한 (end-labeled)올리고뉴클레오타이드 탐침을 사용하여 그의 존재를 탐지하는 것이 어려울 수도 있다. 탐침 탐지 시스템의 민감도(sensitivity)를 증진시키는데에는 상당한 노력이 소요되지만, 통상적으로 이용가능한 방법을 사용하여 용이하게 충분히 탐지 가능한 것이 다량 존재하므로 표적의 서열을 증대시킴에 관한 연구가 거의 이루어지지 않았다.
현존하는 서열로부터 또는 새로이(de novo) 핵산을 합성하는 여러 가지 방법이 문헌에 기술되어 있다. 이들 방법은 완전히 특성을 부여한 서열의 주어진 핵산을 다량으로 제조함을 가능하게 한다.
새로이 핵산을 합성하는 한 알려진 방법에는 뉴클레오사이드(nucleoside) 유도체로부터 핵산을 유기적으로 합성하는 방법이 포함된다. 이 합성법은 용액에서 또는 고체 지지체 상에서 수행할 수 있다. 유기 합성법의 한 형태로 포스포트리에스테르 방법(phosphotriester method)이 있는데, 이 방법은 유전자 단편 또는 짧은 유전자를 제조하는데 이용된다. 포스포트리에스테르 방법에서, 올리고뉴클레오타이드 들은 제조된 다음 함께 연결되어 더 긴 핵산을 형성할 수 있다. 이 방법의 상세한 설명은 하기 문허[Narany S. A., et al.,.,, 90(1979)] 및 미합중국 특허 제4,356,270호에 기술되어 있다. 이 특허에는 소마토스타틴 유전자(someatostatin gene)의 합성 및 클로닝(cloning)에 관한 내용이 기술되어 있다.
유기 합성법의 두 번째 형태로는 포스포디에스테르방법(phosphodiester method)이 있는데, 이 방법은 tRNA 유전자를 제조하는데 이용된다. 이 방법의 상세한 설명은 하기 문헌(Brown. E. L., et al.,.,, 109(1979)]에 기술되어 있다. 포스포트리에스테르 방법에서와 마찬가지로 포스포디에스테르방법에는, 결과적으로 함께 연결되어 목적하는 핵산을 형성하는 올리고클레오타이드의 합성법이 포함된다.
상기의 드 노보 합성법(de novo synthesis)들이 장쇄 핵산의 합성에 이용될 수는 있지만, 이들 방법은 다량의 핵산 합성에 매우 실용적으로 사용되지는 않는다. 상기의 방법은 힘이 들고 시약을 소비하여, 값비싼 장비와 시약을 필요로 하며, 총괄 효율(overall efficiency)이 낮다. 낮은 총괄 효율은 올리고뉴클레오타이드의 합성 및 연결 반응의 비효율에 의해 야기될 수 있다. 긴 핵산을 합성함에 있어, 또는 심지어 더 짧은 핵산을 다량 합성함에 있어, 많은 올리고뉴클레오타이드가 합성되는데 필요하며 많은 연결 반응이 요구되리라고 생각된다. 결과적으로, 이들 방법들은 어떤 목적하는 핵산을 다량 합성하는데에는 실용적이지 못할 것이다.
초기에 존재하는 소량의 핵산으로부터 다량의 핵산을 제조하는 방법들도 또한 있다. 이들 방법에는 적절한 숙주 시스템에서 핵산을 클로닝시키는 방법이 포함되는데, 이 방법에서는 목적하는 핵산은, 숙주를 형질전환시키는데 사용되는 적절한 백터내로 삽입시킨다. 숙주를 배양하는 경우 벡터는 복제되므로, 목적하는 핵산을 더 많은 복제본으로 제조한다. 핵산 단편을 아클로닝시킴에 관한 간단한 설명은 하기 문헌[Maniatis, T.m, et al.,., Cold Spring Harbor Laboratory PP. 390-401(1982)]에 기술되어 있다. 또한 이에 관련된 기술(techniques)은 미합중국 특허 제 4,416,988호 및 제 4,403,036호에 기술되어 있다.
핵산을 합성하는 세 번째 방법은, 미합중국 특허 제 4,293,652호에 기술되어 있으며, 상기 기술한 유기 합성의 하이브리드 및 분자 클로닝 방법이다. 이 방법에서 목적하는 핵산 서열을 구성하는 올리고뉴클레오타이드의 적절한 수는 유기적으로 합성하여, 각각 후속의 삽입과정전에 성장에 의해 증대되는 벡터내로 연속적으로 삽입한다.
본 발명은 분자의 클로닝 방법과 약간 유사한 점을 내포하고 있다 : 하지만, 본 발명은 특정 유기체의 증식(propagation)은 포함하지 않으므로 본 발명에 수반되는 가능한 위험 또는 불편한 점을 피하고 있다. 본 발명은 또한 목적하는 서열에 관련되지 않은 핵산 서열의 합성을 필요로 하지 않으므로, 본 발명은 복잡한 생물학적 혼합물로부터의 생성물의 값비싼 정제 과정의 필요를 미연에 방지한다.
본 발명은 핵산 및 이의 혼합물에 존재하는 하나 이상의 특정한 핵산 서열을 프라이머 및 유도제를 사용하여 증대시키는 방법이다. 1개의 프라이머의 증대 생성물은, 다른 것과 하이브리드화되는 경우, 목적하는 특정의 핵산 서열의 생산을 위한 주형(template)이 되며, 역으로, 과정은 서열의 목적하는 양을 생산하는 데 필요한 만큼 자주 반복할 수 있다. 이 방법은 표적의 서열로부터 핵산을 다량 생산함에 대해 상기 기술한 방법들보다 더욱 효율적이라고 기대되며 비교적 단기간에 그러한 핵산을 생산하리라고 기대된다. 본 발명의 희귀 종 핵산의 효과적인 탐지를 위해 핵산 혼합물중에 존재하는 그러한 희귀 종의 핵산을 증대시키는 데 특히 효과적이다.
더욱 상세히 설명하자면, 본 발명은 하기 단계들로 이루어짐을 특징으로하여, 핵산 또는 핵산의 혼합물을 함유하는 적어도 하나의 특정의 핵산 서열을 증대시키는 방법을 제공하며, 여기에서 각 핵산은 같거나 또는 다른 길이의 두 개의 상보적인 서열로 구성되어 있다.
(a) 각각 다른 특명 서열을 증대시키기 위해, 각 프라이머의 증대 생성물이 각 핵산 본쇄에 상보적으로 합성되는 조건하에서 본쇄들을 2개의 프라이머로 처리하며, 여기에서 상기의 트라이머 또는 프라이머들은 1개의 프라이머로부터 합성된 증대 생성물이, 그의 상보서열(complement)로부터 분리되는 경우, 다른 프라이머의 증대 생성물의 합성을 위해 주형으로 제공될 수 있도록 감각의 특정 서열의 각 본쇄에 실질적으로 상보되도록 선택된다 : (b) 프라이머 증대 생성물을, 그들이 합성된 주형으로부터 분리시켜 일본쇄 분자를 제조하고 : (c) 단계 (b)로부터 생산한 일본쇄 문자를 프라이머 증대 생성물이 단계(b)에서 제조된 일본쇄 각각을 주형으로 사용하여 합성되는 조건하에 단계(a)의 프라이머로 처리한다.
단계들은 연속적으로 또는 동시에 수행한다. 게다가 단계 (b) 및 (c)는 서열이 목적하는 수준까지 증대될 때까지 반복할 수 있다.
본 발명은 현존하는 완전히 특이한 서열의 핵산을 다량으로 제조하는데 뿐만 아니라 있기는 하나 완전히 특이하지 않은 핵산 서열을 제조하는데 있어 유용할 수 있다. 각각의 경우에 증대시킬 서열의 초기 복제본은, 비록 순수하거나 또는 분리된 분자일 필요는 없더라도 수득할 수 있어야만 한다.
본 명세서에서 프라이머, 탐침, 탐지될 올리고머 단편, 올리고머 조절자 및 표지하지 않은 블로킹 올리고머(unlabeled blocking oligomer)에 관하여 사용된 "올리고뉴클레오타이드(oligonucleotide)"는 데옥시리브 뉴클레오타이드 또는 리보뉴클레오타이드 2개 이상, 바람직하게는 3개 이상으로 이루어진 분자로 정의된다. 그의 정확한 크기는 여러 가지의 인자(factors)에 따라 다르며, 이는 올리고뉴클레오타이드의 궁극적인 작용 또는 용도에 따라 다른 것이다.
본 명세서에 사용된 "프라이머(primer)"는 정제된 반응 분해물에서 자연적으로 발생하든 또는 합성적으로 제조하던, 핵산 본쇄에 상보적인 프라이머 증대생성물의 합성이 야기될 수 있는 조건하, 즉 적절한 온도 및 pH에서 뉴클레오타이드, 및 DNA 폴리머라제와 같은 유도체 존재하에 놓여지는 경우 합성의 시발점으로 작용할 수 있는 올리고뉴클레오타이드에 관한 것이다. 프라이머는 증대시 최대 효율을 위해서 바람직하게는 일본쇄이나, 이본쇄일 수도 있다. 이본쇄인 경우, 프라이머는 증대 생성물을 제조하기 위해 사용되기 전에 먼저 그의 본쇄들로 분리시킨다. 바람직하게는, 프라이머는 올리고데옥시리보뉴클레오타이드이다. 프라이머는 유도제 존재하여 증대 생성물의 합성을 준비하기 위해서는 충분히 길어여만 한다. 프라이머들의 정확한 길이는 온도 및 프라이머 원(source of primer)을 포함한, 여러 가지 인자에 따라 다를 것이다. 예를들면, 표적 서열의 복잡성에 따라, 올리고뉴클레오타이드 프라이머는 비록 올리고뉴클레오타이드를 거의 함유하지 않을지라도, 전형적으로 15-25 또는 그 이상의 올리고뉴클레오타이드를 함유한다. 짧은 프라이머 분자는 통상적으로 주형과 함께 충분히 안정한 하이브리드복합물을 형성하기 위해서 더 차가운 온도를 요구한다.
본 명세서에서 프라이머들은 증대시킬 각 특정서열의 상이한 본쇄에 "상당히(substanitally)"상보되도록 선택된다. 이것은 프라이머들이 그들 각자의 본쇄들과 하이트리드화하기 위해서 상보적이어야 한다는 것을 의미한다. 그러므로, 프라이머 서열은 주형의 정확한 서열을 반영할 필요가 없다. 예를들면, 비-상보적은 뉴클레오타이드 단편은, 본쇄에 상보적인 프라이머 서열의 나머지와 함께, 프라이머 5'말단에 결합될 수 있다. 또한 비-상보적인 염기 또는 더 긴 서열을 프라이머내로 산개시킬 수 있는데, 단, 프라이머 서열은 그와 함께 하이브리드화 하기 위해 증대될 본래의 서열과 충분한 상보성을 가지며, 그러므로 다른 프라이머의 증대 생성물의 합성을 위해 주형을 형성한다.
본 명세서에 사용된 바대로, "제한 엔도뉴클레아제(restriction endonucleases)" 및 "제한효소(restrictionenzymes)"는 각각 특성의 뉴틀레오타이드 서열에서 또는 서열 근처에서 이본쇄 DNA를 절단하는 세균효소에 관한 것이다.
본 명세서에 사용된 바대로, "DNA 다형현상(polymorphism)"은 2개, 이상의 상이한 뉴클레오타이드 서열들이 DNA에서 특정 부위에 존재할 수 있는 상태에 관한 것이다.
"제한 단편 길이 다형현상(RFLP)"은 특정의 제한 엔도뉴클레아제로 분해시킴에 의해 형성된 제한 단편들의 길이에서 각각 사이의 차이에 관한 것이다.
본 발명은 직접적으로는 핵산에서 발견되는 하나 이상의 목적으로 특정 핵산 서열을 증대시키는 방법에 관한 것이다. 다량의 특정 서열이 본 발명에 의해 생성될 수 있으므로, 본 발명은 DNA 또는 mRNA의 클로닝효율을 증대시키기 위해, 및 표적서열의 탐지를 용이하게 하기 위해 이 표적서열을 증대시키기 위해 사용될 수 있다.
일반적으로 본 방법은 (a) 요구되는 서열의 말단이 올리고뉴클레오타이드가 합성되어 그들과 하이브리드화될 정도로 충분히 공지되고, (b) 소량의 서열이 연쇄 반응을 개시할 수 있도록 주어진, 1개 이상의 특정핵산 서열을 반응 단계에 포함된 수와 비교해서 지수적인 양으로 제조하기 위한 연쇄 반응을 포함한다. 연쇄 반응의 산물은 사용한 특정 프라이머의 말단에 상응하는 말단을 가진 분리된 핵산 이본쇄일 것이다.
어느 핵산원이건 정체되거나 또는 정제되지 않은 상태로 출발 핵산 또는 핵산들로서 사용하여, 목적하는 특정 핵산 서열을 함유하게 하거나 아마 함유하게 할 수 있다. 따라서 본 발명은, 예를들면 일본쇄 또는 이본쇄 일 수 있는 DNA 똔느 RNA와 전령 RNA를 사용할 수 있다. 게다가 각각의 일본쇄를 함유하는 DNA-RNA 하이브리드를 사용할 수 있다. 이들 핵산의 어느 혼합물도 사용할 수 있으며, 또는 본문에서 동일 또는 상이한 프라이머를 사용하는 선행 증대 반응으로부터 제조한 핵산을 사용할 수도 있다. 증대시킬 특정 핵산 서열은 단지 큰 분자의 단편일 수 있거나, 또는 초기에 분리된 분자로서 존재할 수 있어 특정 서열의 전체 핵산을 구성하고 있다. 증대될 서열이 초기에 순수한 형태로 존재할 필요는 없고 : 유기체가 특정생물학적 샘플의 단지 아주 작은 부분을 구성하는 특정 미생물로 인해 핵산 서열의 일부 또는 전 인체 DNA에 포함된 β-글로빈 유전자 부분과 같이, 복합 혼합물의 소부분일 수 있다. 출발 핵산은 동일 또는 상이한 목적하는 특정 핵산 서열을 하나이상 함유할 수 있다. 그러므로 본 발명은 1개의 특정 핵산 서열을 다량으로 제조하는데 뿐만 아니라 동일 또는 상이한 핵산 분자상에 위치한 상이한 특정핵산 서열을 1개 이상 동시에 증대시키는데 유용하다.
핵산 또는 핵산들은 어느 원천, 예를들면, pBR 322와 같은 플라스미드, 클론된 DNA 또는 RNA 또는 세균, 효모, 비루스 및 식물 또는 동물과 같은 고등생물을 포함한 어떤 원천의 자연 DNA 또는 RNA로부터 수득할 수 있다. NDA 또는 RNA는 혈액, 융몸모막의 융모(chorionic villi) 또는 양수 세포(ammiotic cells)와 같은 조직 물질로부터 하기 문헌[Maniatis et al., Molecular Cllning(1982), 280-281]에 기술되어 있는 여러 가지 기술에 의해 추출할 수 있다.
어느 특정 핵산 서열도 본 발명에 의해 제조할 수 있다. 단지 서열의 양말단에 있는 충분한 수의 염기가 상세히 알려져 있어 2개의 올리고뉴클레오타이드 프라이머를 목적하는 서열의 다른 분쇄에 서열중의 상대위치에서 하이브리드화하도록 제조할 수 있고, 1개의 프라이머로부터 합성된 증대 생성물이 그의 주형(상보서열)으로부터 분리될 경우, 한정된 길이의 핵산으로 기타의 프라이머를 증대시키기 위한 주형으로 기여할 수 있으면 된다. 서열의 양단면에 있는 염기에 관한 지식이 많으면 많을수록 표적핵산 서열용 프라이머의 특이성이 크고, 따라서 본 발명의 효율도 높아진다. 이 후 사용되는 단어, 프라이머는 특히 증대시킬 단편의 종료 서열(들)에 관한 정보에서 다소 모호한 경우 프라이머 1개 이상을 가리킨다. 예를들면, 단백질 서열 정보로부터 핵산 서열을 추론하는 경우 유전 코드의 축퇴성(degeneracy)을 기본으로 하여 모든 가능한 코돈 변이형을 나타내는 서열을 함유하는 프라이머의 집합이 각 본쇄에 사용될 것이다. 이 집합으로부터의 1개 프라이머는 증대시킬 목적 서열의 말단과 100% 동일할 것이다.
올리고뉴클레오타이드 프라이머는 어느 적합한 방법 예를들면, 상술한 포스토트리에스테르 및 포스포디에스테르방법, 또는 그의 자동조작되는 태양과 같은 방법을 사용하여 제조할 수 있다. 그와 같이 자동조작되는 한 태양에서 디에틸 포스포아미디트가 출발 물질로서 사용되며 하기 문헌[Beaucage et al.,(1981), 22 : 1859 - 1862]에 기술된 바대로 합성할 수 있다. 개질된 고체 지지대상에서 올리고뉴클레오타이드를 합성하는 한 방법이 미합중국 특허 제4,458,067호에 기술되어 있다. 생물학적 원천으로부터 분리한 프라이머(제한 엔도뉴클레아제와 같은)를 사용할 수도 있다.
특정 핵산 서열은 그 서열을 함유하는 핵산을 주형으로 사용해서 제조한다. 만일 핵산이 이본쇄를 함유하는 경우, 주형으로 사용하기 전에 프라이머 증대 생성물의 합성과 분리단계로서 또는 동시에 핵산의 본쇄를 분리시킬 필요가 있다. 이 본쇄 분리는 물리적, 화학적 또는 효소적 방법을 포함하는 적당한 방법으로 수행할 수 있다. 핵산의 본쇄를 분리시키는 한가지 물리적 방법이 핵산이 완전히(>99%) 변성될 때까지 가열시키는 것이다. 전형적인 열 변성은 약 80 내지 105℃ 범위의 온도에서 약 1 내지 10분 범위의 시간이다. 본쇄 분리는 또한 헬리카제(helicase) 활성을 갖고 리보 ATP의 존재하에서 DNA를 변성시키는 것으로 알려져 있는 RecA 또는 헬리카제로서 공지되어 있는 효소군으로부터의 효소에 의해 효소할 수 있다. 헬리카제로 핵산의 본쇄를 분리하는데 적합한 반응 조건은 하기 문헌[kuhn Hoffmann-Berling, , 43 : 63 (1978)]에 기술되어 있고, RecA를 사용하는 기술은 하기 문헌[C.Radding,: 405-37(1982)]에 시사되어 있다.
증대시킬 서열은 함유하는 원형 핵산이 일본쇄인 경우, 그의 상보서열은 거기에 1 또는 2개의 올리고뉴클레오타이드 프라이머를 가하여 합성한다. 적절한 단일 프라이머를 가하면, 프라이머 증대 생성물이 프라이머, 합성의 유도체 또는 촉매 및 하기에 기술된 4가지 뉴클레오타이드의 존재하에 합성된다. 생성물은 일본쇄 핵산에 부분적으로 상보적이며, 핵산 본쇄와 하이브리드화하여 상이한 길이의 이본쇄를 형성하고, 이어서 상술한 바와같이 일본쇄로 분리되어 2개의 분리된 상보적 일본쇄를 산출할 것이다. 또한 2개의 적당한 프라이머를 일본쇄 핵산에 가하여 반응을 수행할 수도 있다.
원형 핵산이 증대시킬 서열을 구성하고 있는 경우, 생성된 프라이머 증대 생성물(들)은 원형 핵산의 본쇄에 완전히 상보적이어서 서로 하이브리드화되며 일본쇄 분자로 분리될 동일한 길이의 이본쇄를 형성할 것이다.
핵산이 본래 이본쇄이거나 일본쇄이거나 핵산 또는 핵산들의 상보적인 본쇄들이 분리되면, 본쇄는 부가적인 핵산 본래의 합성을 위한 주형으로서 사용하기가 용이하다. 이 합성은 적당한 방법을 사용하여 수행한다. 통상 합성은 바람직하게는 pH 7-9 더욱 바람직하게는 약 8인 완충 수용액에서 수행한다. 바람직하게는 2개의 올리고뉴클레오타이드 프라이머의 물과량(클론된 핵산에 대해서는 통상 프라이머 : 주형이 약 1000 : 1, 게놀의 핵산에 대해서는 통상 프라이머 : 주형이 약 106: 1)을 분리된 주형 본쇄가 함유된 완충액에 가한다. 그러나 본 발명을 감별 적용에 사용하는 경우, 상보적인 본쇄의 양을 알 수가 없어 상보적인 본쇄의 양에 대한 프라이머의 양을 정확히 측정할 수 없다. 하지만 실용적인 문제로서 통상 가해지는 프라이머의 양은 증대시킬 서열이 복잡한 장쇄 핵산 본쇄의 혼합물에 함유되어 있는 경우 상보적인 본쇄(주형)의 몰과량일 것이다. 큰 물과량은 큰 몰과량은 방법의 효율을 증진시키는데 바람직하다.
데옥시리보뉴클레오타이드 트리포스페이트 dATP, dCTP, dGTP 및 TTP도 적량을 합성 혼합물에 가하고, 생성된 용액을 약 90 내지 100℃로 약 1내지 10분, 바람직하게는 1 내지 4분동안 가열한다. 이 가열기간이후 용액을 플라이머 하이브리드화에 바람직한 실온으로 냉각한다. 냉각된 혼합물에 프라이머 증대 반응을 유도 또는 촉매(여기서는 "유도체"라 칭함)하는 적당한 제제를 가하고, 당분야에 공지된 조건하에 반응을 수행한다. 이 합성 반응은 실온 내지 유도제가 더 이상 효율적으로 반응하지 못하는 온도 이상까지에서 수행할 수 있다. 따라서 예를들면, DNA 폴리머라제를 유도제로 사용한 경우 온도는 통상 약 40℃보다 높지 않다. 가장 편리하게는 실온에서 반응시킨다.
유도제는 효소를 포함하여 프라이머 증대 생성물의 합성을 수행하는 특성 화합물 또는 시스템일 수 있다. 이 목적에 적합한 효소는, 예를들면,DNA 폴리머라제, I.DNA 폴리머라제 I의 클레노우 단편, T4DNA 폴리머라제, 기타 가능한 DNA 폴리머라제, 역전사 효소 및 내열성 효소를 포함한여 적당한 방식으로 뉴클레오타이드를 배합하여 각 핵산 본쇄에 상보적인 프라이머 증대 생성물을 형성하는 기타 효소 등이다. 일반적으로 합성은 각 프라이머의 3'말단에서 개시하여 합성이 종료될 때까지 주형 본쇄를 따라 5' 방향으로 진행하며, 상이한 길이의 분자를 제조한다. 그러나 상술한 동일 방법을 사용하여 5'말단에서 합성을 개시하고 다른 방향으로 진행하는 유도제도 있을 수 있다.
새로 합성된 본쇄 및 그에 상보적인 핵산 본쇄는 이본쇄 분자를 형성하여 본 발명의 연속 단계에 사용된다. 다음 단계에서 이본쇄 분자의 본쇄는 상술한 특정 방법을 사용 분리하여 일본쇄 분자를 제공한다.
새로운 핵산은 일본쇄 분자상에서 합성된다. 반응을 진행시키는데 필요하면 부가적인 유도제, 뉴클레오타이드 및 프라이머를 상술한 조건하에 첨가할 수 있다. 역시 합성은 올리고뉴클레오타이드 프라이머의 한쪽 말단에서 개시하고 주형의 일본쇄를 따라 진행하여 부가적인 핵산을 생성한다. 이 단계후, 증대 생성물의 반이 두 프라이머에 의해 결합된 특정핵산 서열을 구성한다.
본쇄 분리 및 증대 생성물 합성의 단계는 특성 핵산 서열의 목적하는 양을 생산하기 위해 필요한 만큼 자주 반복할 수 있다. 계속 하기에 상세히 설명하겠지만, 생성된 특정 핵산 서열의 양은 지수적 양상으로 축적된다.
첫 번째 핵산 또는 핵산의 혼합물로부터 한가지 이상의 특정 핵산 서열을 제조하려고 할 때는 상이한 올리고뉴클레오타이드 프라이머의 적정한 수를 사용한다. 예를들면 2개의 상이한 특정 핵산 서열을 제조할 경우 4개의 프아미어가 사용된다. 프라이머중 2개는 특정 핵산 서열의 하나에 대해 특이하고, 다른 2개의 프라이머는 두 번째 특정 핵산 서열에 대해 특이하다. 이런 식으로 2개의 상이한 특정 서열의 각각을 본 발명에 의해 지수적으로 생산할 수 있다.
본 발명은 각 단계후에 새로운 시약을 가하거나 동시에 가하는 방법, 모든 시약을 개시 단계에 가하거나, 부분씩 단계적으로 및 부분씩 동시에 가하는 방법, 모든 시약을 개시 단계에 가하거나, 부분씩 단계적으로 및 부분씩 동시에 가하는 방법, 신선한 시약을 주어진 모든 단계후에 가하는 방법등 단계적 방법으로 수행할 수 있다.
만약 열에 약한 효소의 경우에, 열과 같은 분쇄 분리 방법을 사용하여 유도체를 불활성화시킨다면, 이어서 매 분쇄 분리 단계 이후 유도제를 보충시킬 필요가 있다. 동시 방법은 효소적 수단을 본래 분리 단계에 이용한 경우 사용할 수 있다. 동시적 방법에서 반응 혼합물은 목적하는 서열을 함유하는 핵산 본쇄(들)의 예, 분쇄-분리 효소(예 : 헬리카제), ATP와 같이 적당한 본쇄-분리효소용 에너지원, 4개의 뉴클레오타이드, 과량의 올리고뉴클레오타이드 프라이머, 및 예를들어DNA 폴리머라제 1의 클로노우 단편과 같은 유도제를 함유한다. 동시적 방법에서 변성시 열을 사용한 경우, 열안정성 폴리미라제와 같은 내열과 같은 유도제를 함유한다. 동시적 방법에서 변성시 열을 사용한 경우, 열안정성 폴리머라제와 같은 내열성 유도제를 사용하여 상승온도에서, 바람직하게는 유도제에 따라 65-90℃에서 조작하며, 이 온도에서 핵산은 일본쇄와 이본쇄를 동등하게 구성한다. 길이가 짧은 핵산을 위해서는 약 50℃의 낮은 온도를 사용한다. 그 이상의 온도는 효소가 분해하는 온도 또는 프라이머 하이브리드화가 불충분한 수준으로 일어나는 온도 이상에 의존한다. 그런 내열성 효소는 예를들면 하기 문헌[A. S. Kaledin et al,, 644-651(1980)]에 기술되어 있다. 방법의 각 단계는 개시에서의 모든 시약의 존재에도 불구하고 연속해서 일어난다. 가외의 물질을 필요에 따라 가할 수 있다. 적당한 시간이 경과하여 특정 핵산 서열의 목적하는 양이 생성된 후, 공지된 방법으로 효소를 불활성화하거나 반응의 성분을 분리함으로써 반응을 정지시킬 수 있다.
본 발명의 방법은 계속적으로 수행할 수 있다. 자동조작되는 방법의 한 태양에서, 반응은 변성 부위, 시약 첨가 부위, 반응 부위를 통해 순환한다. 다른 태양에서 프라이머 증대 생성물의 합성시 사용되는 효소를 컬럼에 고정시킬 수 있다. 기타 반응 성분은 펌프에 의해서 컬럼과 일련의 가열 코일을 통해 순환되며 따라서 생성된 핵산이 효소의 불활성화 없이 반복해서 변성될 수 있다.
본 발명은 상보적인 본쇄 [S+] 및 [S-]으로 구성된 목적 서열[S]를 함유하는 이본쇄 DNA를 핵산으로 사용하여 하기에 도식적으로 설명되고 있다. 첫 번째 및 매 후속적 반응 사이클도중 원형 주형상에 각 올리고뉴클레오타이드 프라이머의 증대는 단지 1개의 프라이머로 종료하는 무한한 걸이의 새로운 ssDNA분자 생성물을 생성할 것이다. 이후 생성물은 "긴(long) 생성물"이라 칭하며, 선상으로 축적할 것이다 : 즉 어느 사이클수 이후에 존재하는 양은 사이클 수에 비례한다.
생성된 긴 생성물 후속 사이클중에서 하나의 또는 다른 올리고뉴클레오타이드 프라이머용 주형으로 작용하여 목적하는 서열 [S+] 또는 [S-]분자를 생성한다. 이 분자들은 또한 하나의 또는 다른 올리고뉴클레오타이드 프라이머용 주형으로 작용하여 그밖의 [S+] 및 [S-]를 생성하고, 다라서 연쇄 반응이 유지되며, 결과적으로, 사이클의 수화 비교해서 지수적 비율로 [S]의 축적을 초래한다.
의도한 것들외에 올리고뉴클레오타이드 하이브리드화에 의해 형성된 부산물은 자기-촉매적이지 못하며 (소수의 경우는 제외) 따라서 선상 비율로 축적한다.
증대시킬 특정 서열, [S]는 다음과 같이 도식적으로 인용할 수 있다 :
[S+] 5' AAAAAAAAAAXXXXXXXXXXCCCCCCCCCC 3'
[S-] 3' TTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
적당한 올리고뉴클레오타이드 프라이머는 다음과 같다.
프라이머 1 : GGGGGGGGGG
프라이머 2 : AAAAAAAAAA
그래서 만일 [S]를 함유하는 DNA
.....zzzzzzzzzzzzzzzAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzzzz.....
.....zzzzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGGzzzzzzzzzzzzzzzz.....
이 일본쇄로 분리되고 그의 일본쇄가 프라이머 1과 2에 하이브리드화되면, 하기 증대 반응이 4개의 데옥시리보 뉴클레오사이드 트리포스페이트의 존재하여 DNA 폴리머라제에 의해 촉매될 수 있다 :
.....zzzzzzzzzzzzzzzzAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzzzzz......
원형 주형 본체+
원형 주형 본체-
.....zzzzzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGGzzzzzzzzzzzzzzzzz.....
형성된 2개의 이본쇄의 변성시, 생성물은 다음과 같다 :
3'.....zzzzzzzzzzzzzzTTTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
새로 합성된 긴 생성물 1
5'.....zzzzzzzzzzzzzzAAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzz.....3'
원형 주형 본체+
3'.....zzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGGzzzzzzzzzzzzzz.....5'
원형 주형 본체-
5'AAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzz...3'
새로 합성된 긴 생성물 2
이들 4개의 본쇄를 다음 사이클에서 프라이머 1 및 2와 하이브리드화시킬 경우, 유도체는 하기 반응을 촉매한다.
3'.....zzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
새로 합성된 긴 생성물 1
5'.....zzzzzzzzzzzzzzAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzz..... 3'
원형 주형 본체+
3'......zzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGGzzzzzzzzzzzzzz..... 5'
원형 주형 본쇄+
5'AAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzz...... 3'
상기 4개의 이본쇄가 분리되면, 하기 본쇄가 된다 :
5' AAAAAAAAAAXXXXXXXXXXCCCCCCCCCC 3'
새로 합성된 [S+]
3'......zzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
첫번째 사이클에서 합성된 생성물 1
3'......zzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
새로 합성된 긴 생성물 1
5'......zzzzzzzzzzzzzzzzAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzz.....3'
원형 주형 본쇄+
5'AAAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzz......3'
새로 합성된 긴 생성물 2
3'zzzzzzzzzzzzzzTTTTTTTTTTYYYYYYYYYYGGGGGGGGGGzzzzzzzzzzzzzz......5'
원형 주형 본쇄+
3'TTTTTTTTTTYYYYYYYYYYGGGGGGGGGG 5'
새로 합성된 [S-]
5'AAAAAAAAAAXXXXXXXXXXCCCCCCCCCCzzzzzzzzzzzzzzzz......3'
첫 번째 사이클에서 합성된 긴 생성물 2
한 개 프라이머의 올리고뉴클레오타이드 서열 및 다른 상보적 서열로 종결되는 각 본쇄는 생산하기에 바람직한 특정 핵산 서열임을 알 수 있다.
이 방법의 단계는 존재하는 프라이머 1 및 2. 유도체 및 뉴클레오타이드의 양에 의해 제한받을 뿐, 무한하게 반복할 수 있다. 원형 핵산의 양은 전 방법에서 일정하게 남아있는데 이것은 복제되지 않기 때문이다. 긴 생성물의 양은 그들이 단지 원형 핵산으로부터 제조되기 때문에 일직선으로 증가한다. 특정 서열의 양은 지수적으로 증가한다. 따라서 특정 서열은 우세한 종으로 된다. 이것은 하기 표에 설명되어 있는데, 표는 각 사이클에서의 효율을 100%로 가정하여 n 사이클 후 이론적으로 존재하는 본 종들의 상대량을 가리킨다 :
[0내지 n 사이클 후]
[이본쇄의 수]
일본쇄 핵산을 주형으로 사용하면, 사이클당 단지 하나의 긴 생성물이 형성된다.
여기에서의 방법은 적당한 발현 벡터속에 삽입시 특정 핵산 서열을 클론하는데 사용할 수 있다. 이어서 벡터는 적당한 숙주 유기체를 형질전환시키는데 사용하여 재조합 DNA 기술의 표준 방법에 의해 서열의 유전산물을 생산할 수 있다.
게다가 본 방법은 생체의 돌연변이시 사용할 수 있다. 올리고데옥시리보뉴클레오타이드 프라이머가 증대시킬 DNA서열에 정확히 상보적일 필요는 없다. 그들이 증대될 서열과 폴리머라제 효소 또는 사용한 다른 유도제에 의해 충분히 하이브리드화할 수 있으면 된다. 사용한 프라이머가 원혀 주형에 완전히 상보적이 아닌 폴리머라제 연쇄 반응의 생성물은 주형보다는 오히려 프라이머 서열을 함유해서 생체외 돌연변이를 유발할 것이다. 다음 사이클에서 이 돌연변이는 저하하지 않는 효율로 증대되는데, 이유는 더 이상 잘못 짝지어진 프라이밍(primings)이 요구되지 않기 때문이다. 그렇게 생성된 돌연변이체는 표준 분자 생물학적 기술에 의해 적당한 벡터로 삽입하며, 이 벡터상에 변화단백질의 생성 능력과 같은 돌연변이 특성을 부여할 수 있다.
상술한 바와 같이 변화된 DNA서열을 만드는 방법은 상이한 프라이머를 사용하여 변화된 DNA상에서 반복함으로써 그밖의 서열 변화를 유도할 수 있다. 이런 식으로 일련의 변이 서열이 점차 생성되며, 이 일련에 대한 새로운 부가는 바로 전의 것과 조금씩 다를 수 있으나 원형의 DNA원천 서열과는 점차적으로 크게 다르다. 이렇게 하여 궁극적으로 변화시킬 수 있으며, 아주 심하게 잘못연결된 프라이머는 기능을 갖지 못하므로 1단계로서는 불가능하다.
더불어 프라이머는 프라이머의 충분한 양이 증대시킬 서열에 상보적인 서열을 함유하는 비-상보적 서열을 그의 서열의 일부로서 함유할 수 있다. 예를 들면 주형 서열에 상보적이 아닌 뉴클레오타이드 서열(예 : 프로모터, 링커, 코드화서열 등)은 프라이머의 양쪽 또는 한쪽의 5'말단에 부착될 수 있으며, 그로인해 증대 공정의 생성물에 덧붙일 수 있다. 증대 프라이머를 가한 후, 충분한 사이클을 진행시켜 비-상보적인 뉴클레오타이드 삽입물을 함유하는 신규 주형의 목적하는 양을 취할 수 있다. 이것은 단순한 기술을 사용하여 비교적 짧은 시간(예 : 2시간 또는 그 이하)에 다량의 결합 단편을 생산하게 한다.
이 방법은 또한 감염성 질병, 암과 같은 유전자성 질환 또는 세포성 질환과 결부된 특정 핵산 서열의 검출 및/또는 특정화를 하는데 사용할 수 있다. 증대는 분석할 핵산이 매우 작은 경우, 예를 들면 태아 세포로부터 수득한 DNA를 사용하여 겸상적혈구 빈혈증의 태아가 진단시 유용하다. 증대는 그와 같은 분석을 비-방사성 검출법을 사용하여 본래 둔감한 소량의 샘플에 대해 수행할 때, 또는 방사성 기술을 사용하되 신속한 검출이 요구될 대 특히 유용하다.
본 발명의 목적인 유전병은 특정 유기체로부터 게놈 DNA상의 특정 결실 및/또는 돌연변이, 예를 들면, 겸상적혈구 빈혈, 낭포성 섬유화(cystic fibrosis), α-중증성 지중해 빈혈(thalessemia), β-중증성 지중해 빈혈등을 포함한다. 겸상적혈구 빈혈은 본 방법에 의해서 적당한 DNA 서열의 증대에 이어 RFLP-형분석 또는 올리고머 제한 분석을 통해 쉽게 검출할 수 있다. α-중증성 지중해 빈혈은 서열의 부재에 의해 검출할 수 있고, β-중증성 지중해 빈혈은 질병을 일으키는 돌연변이에 근접하여 연결된 다형성 제한 부위의 존재에 의해 검출할 수 있다.
이 모든 유전적 질병은 적당한 서열을 증대시키고 방사성 탐침 사용없이 서던 블롯(Southern blots)으로 그것을 분석하여 탐지할 수 있다. 그런 방법에서, 예를 들면 목적하는 아주 낮은 정도의 서열을 함유하는 양수로부터 소량의 DNA 샘플을 증대시키고, 제한 효소로 절단하며, 서던 블롯 기술을 통해 분석한다. 비-방사성 프로브의 사용은 고수준의 증대 신호에 의해 용이하게 한다.
또 다른 태양에서 작은 DNA 샘플은 통상의 수준으로 증대되고, 쉽게 탐지될 수 있는 뉴클레오타이드유도체(예 :32P-표지 도는 비오틴 표지된 뉴클레오타이드 삼인산)를 최종 DNA 생성물에 직접 결합시킴으로써 증대반응의 그밖의 사이클을 수행하여, 제한 및 전기영동 분리 또는 기타의 적당한 방법으로 분석한다. 이 기술의 실시예가 모델 시스템으로 제5도에 설명되어 있다.
제3도에 모델 시스템으로 설명되고 있는 그밖의 태양에서, 핵산은 증대되기 전에 특정 제한 엔도뉴클레아제에 노출된다. 절단된 서열은 증대될 수 없기 때문에, 증대 단편의 출현은 DNA 샘플의 앞서의 제한에도 불구하고 증대된 서열내에 엔도뉴클레아제를 위한 부위가 부재한다. 증대 서열의 존재 또는 부재는 적당한 방법으로 탐지한다.
이 기술의 실제적 응용은 아래 본문 및 문헌 [R. Saiki et al., Bio Technology, 3 : 1008-1012 (1985)]에 기술된 올리고머 제한 기술을 통해 겸상적혈구 빈혈의 탐지를 용이하게 하는데 있어서의 그의 용도에 의해 설명할 수 있다. 겸상적혈구 빈혈은 β-글로빈 유전자의 6번째 코돈에 단일 염기쌍 변화가 일어나 유발되는 헤모글로빈 질환이다. 제6도는 그의 다형성 부위에서 정상 및 겸상적혈구 β-글로빈 유전자의 서열을 명시하고 있는데, 단선(Single bars)은 경상 유전자에만 존재하는 Dde I 부위의 위치를 표시하고, 복선(double bars)은 비-다형성이고 따라서 정상 및 겸상적혈구 대립인자에 존재하는 Hinf I 부위의 위치를 표시한다. 제7도는 제한 부위 및 별표로 나타낸 표지를 함유할 프로브를 사용하여 정상 β-글로빈 DNA를 올리고머 제한하는 방법을 설명한다. 여기에서 제공된 증대 DNA를 변성시키고 표지 프로브에 아닌(anneal)시킨다. 효소 Dde I는 DNA를 재형성된 Dde I 부위에서 절단하여 표지된 옥타머(Octamer)를 생성한다. 시험에 사용한 조건하에서 옥타머는 이본쇄로부터 분리될 정도로 충분히 작다. 효소 Hinf I의 후속적 부가는 새로운 일본쇄 옥타머에 효과가 없다. 제8도는 β-글로빈 DNA의 겸상적혈구 대립인자에 적용된 동일 방법을 설명하고 있다. 효소 Dde I는 잘못 짝지어진 A-A 염기쌍으로 인해 증대 DNA 및 표지된 프로브에 의해 형성된 이본쇄를 절단하지 못한다. 그러나 효소 Hinf I은 하이브리드를 제한하여 표지된 트리머(trimer)를 생성시킨다. 실제 이 방법은 특정 신호가 각 대립인자의 존재와 결부되기 때문에 야생형에 동형접합성인 것, 겸상형에 동형접합성인 것, 또는 겸상 적혈구 특성에 이형 접합성인 담체로서 개개의를 감별할 수 있다. 상술한 적절한 서열의 증대 방법의 사용은 단지 단일 DNA 표지된 탐침을 이용해서 단일32P 복제 유전자의 신속한 분석을 허용한다.
여러 가지 감염성 질병은 원인 미생물에 특징적인 특정 DNA 서열의 임상적 샘플의 존재에 의해서 진단할 수 있다. 이들은 살모넬라, 클라미디아, 네이세리아(Neisseria)와 같은 세균, 간염 비루스와 같은 비루스 및 말라리아의 원인인 프라스모듐(plasmodium)과 같은 원생동물의 기생충 등이다. 팔코우(Falkow)가 출원한 미합중국 특허 제4,358,535는 감염성 질병의 진단용으로 특정 DNA 하이브리드화 프로브의 용도를 기술하고 있다. 팔코우 방법에 수반되는 문제는 비교적 소수의 독성 유기체가 감염된 환자로부터 임상 샘플에 존재하고 이들로부터 추출한 DNA가 샘플의 총 DNA중 극히 소부분을 구성할 수 있다는 것이다. DNA 샘플의 고정화 및 하이브리드화 검출전에 예상되는 서열의 특수한 증대는 이 방법의 감도 및 특성을 크게 증진시킬 수 있다.
감염성 질병의 진단시 DNA 프로브의 일반적 임상 용도는 만일 워드(ward)의 EP 63,879에 기술된 바와 같이 비-방사성적으로 표지된 프로브를 사용하는 경우 상당히 단순해진다. 이 방법에서는 비오틴-함유 DNA프로브를 아비딘 또는 비오틴-특이성 항체에 연결된 색원성 효소에 의해 검출한다. 이런 형의 검출이 편리하나, 비교적 민감치 못하다. 본 방법에 의한 특정 DNA 증대 및 안정하게 표지된 프로브의 배합은 일반적인 임상 사용에 팔코우 및 워드 방법을 유용하게 만드는데 요구되는 편리함과 민감함을 제공한다.
증대 방법은 또한 단일 복제 사람 유전자로부터 DNA를 충분한 양으로 제조하는데 사용하여 에티디움 브로마이드와 같이 간단한 비-특이성 DNA 염료를 이용하여 직접 DNA 진단을 할 수도 있다.
유기체의 게놈에서 감염성 질병 및 독성적 비정상을 탐지하는 외에, 본 방법은 특정 독성적 상태와 연관되지 않은 DNA 다형현상을 검출하는데 사용할 수도 있다.
하기 실시예들은 설명의 방법으로 제공된 것으로 어떻게 해서든 본 발명을 제한하지는 않는다. 이들 실시예에서 모든 백분율은 고체의 경우 중량에 의한 것이고 액체인 경우 용적에 의한 것이며, 모든 온도는 다른 주해가 없는 한 섭씨 온도이다.
[실시예 1]
ATCC로부터 입수 가능한 pBR322의 47염기쌍 Fok I 제한 단편 상에 함유되어 있는 뉴클레오타이드 서열
5' CCTCGGCACCGTCACCCTGGATGCT 3'
3' GGAGCCGTGGCAGGGACCTACGA 5'
을 갖는 25염기쌍 서열은 하기와 같이 제조한다. 47-bp 단편을 함유하는 pBR 322의 Fok I 분해물은 제공자인 뉴 잉글랜드 바이오랩스 인코포레이티드(New England Biolabs Inc.)사에 의해 제안된 조건에 따라 pBR 322의 Fok I으로 분해시켜 제조한다. 이용되는 프라이머들은 5'd(CCTCGGCACCG)3' 및 5'd(AGCATCCAGGGTG)3'이며, 통상적인 기술을 사용하여 제조한다. 하기 성분들 [즉 상기 기술한 프라이머들 각각 2433피코몰, pBR 322의 Fok I 분해물, 2.4피코몰, dATP 12나노몰, dCTP 22나노물, dGTP 19나노몰 및 TTP 10나노몰]을 25mM 인산칼륨, 10mM 염화마그네슘 및 100mM 염화나트늄으로 이루어진 완충액(pH 7.5) 33㎕에 가한다.
혼합물을 85℃로 5분동안 가열한 다음 주변 온도로 냉각시킨다. 이. 콜라이 DNA 폴리머라제(I)의 클레노우 단편 5-단위를 가하여 온도를 15분 동안 그대로 유지시킨다. 그후, 혼합물을 다시 85℃로 5분동안 가열한 다음 냉각시킨다. 클레노우 단편 5단위를 다시 가하여 반응을 15분동안 수행한다. 가열, 냉각 및 합성단계를 11회 이상 반복 수행한다.
최종 반복 수행후, 분취량 5㎕을 반응 혼합물로부터 취한다. 이것을 85℃로 3분동안 가열한 다음 주변온도로 냉각시킨다. α-P32-데옥시시티딘 트리포스페이트 12.5피코몰 및 클레노우 단편 5단위를 가하여 반응을 15분동안 게속 수행한다. 표지한 생성물을 폴리아크릴아미드겔 전기영동에 의해 검사한다. Fok I 분해물을 유사한 형태로 표지하여 대조용 및 분자량 표지물로 제공한다. 사이클을 13회 반복 수행한 후 확실시 관찰할 수 있는 표지 밴드가 목적하는 25염기쌍 서열이다.
[실시예 2]
목적하는 증대시킬 서열은 인체의 베타-글로빈 유전자내에 함유되어 있으며 겸상적혈구 빈혈증에 관련된 Mst II 부위를 함유(spanning)하는 94염기쌍 서열이다. 이 서열은 제1도에 나타낸 뉴클레오타이드 서열을 갖는다.
[I. 프라이머 합성]
하기 2개의 올리고 데옥시리보뉴클레오타이드 프라이머를 하기 기술한 방법으로 제조한다.
5' CACAGGGCAGTAACG 3' 프라이머 A
5' TTTGCTTCTGACACA 3' 프라이머 B
자동조작 합성방법 : 하기 문헌[Beauage 및 Caruthers : Tetrahedron Letters (1981) 22 : 1859-1862]에 기술된 방법에 따라 합성한 디에틸 포스포르아미드트 들을 바이오서치(Biosearch) SAM-1을 사용하여 뉴클레오사이드 유도된 조절 다공성 유리 지지체에 연쇄적으로 축합시킨다. 이 방법에는 디클로로메탄중에서 트리클로로아세트산을 사용하여 탈트리틸화시키는 반응(detritylation), 활성 양자 공여체로 벤조트리아졸을 사용하는 축합반응, 및 테트라하이드로푸란 및 피리딘중에서 무수 아세트산 및 디메틸아미노피리딘을 사용하여 캐핑시키는 반응(capping)이 포함된다. 사이클을 1회 수행하는 시간은 약 30분이다. 각 단계에서의 수율은 필요한 양이며, 이것은 탈트리틸화 반응 동안에 방출된 디메톡시트리틸 알콜을 회수하여 분광 분석하여 측정한다.
올리고데옥시리보뉴클레오타이드 보호해제 및 정제 방법 : 고체 지지체를 칼럼으로부터 제거하여 밀폐 튜브 내 실온에서 4시간동안 농 수산화암모늄 1㎖에 노출시킨다. 지지체를 여과하여 제거한 다음 부분적으로 보호된 올리고데옥시튜클레오타이드를 함유하는 용액을 5시간동안 55℃로 되게 한다. 암모니아를 제거하고 잔류물을 예비 폴리아크릴아미드 겔에 올려놓는다. 30Volts/㎝에서 90분동안 전기 영동시킨 다음 생성물을 함유하는 밴드를 형광판의 UV 사진으로 확인한다. 밴드를 절단하여 4℃에서 밤새증류수 1㎖로 용출시킨다. 이 용액을 Altech RP/8칼럼에 올려놓고 1% 암모늄 아세테이트 원출액(pH 6.0)중의 아세토니트릴 7 내지 13%로 용출시킨다. 용출액을 260mm에서 UV 흡광도로 모니터(monitor)하고 적절한 분획을 회수하여 고정 용적에서 UV흡광도로 정량 분석하고 진공 원심분리기내 실온에서 증발건고시킨다.
올리고데옥시리보뉴클레오타이드 특징화 : 정제된 올리고뉴클레오타이드 테스트 분취량을 폴리뉴클레오타이드 키나제 및 r-32p-ATP를 사용하여32로 표지한다. 표지한 화합물을 50Volts/㎝에서 45분동안 전기영동시킨 당므 14 내지 20% 폴리-아크릴아미드겔의 사진으로 검사한다. 이 방법으로 분자량을 입증한다. 염기 조성물은, 독액 디에스터라제(venom diesterase) 및 세균성 알칼리 포스파타제를 사용하여 올리고데옥시리보뉴클레오타이드 뉴클레오사이드들로 분해시킨 다음, 분리하고, 역상 HPLC 칼럼 및 10% 아세토니트릴, 1% 암모늄 아세테이트 유동상을 사용하여, 유도된 뉴클레오사이드들을 정량분석하여 측정한다.
[II. DNa 원]
A. 전인체의 야생형 DNA의 추출
정상적인 β-글로빈에 동형 접항성인 인체게놈의 DNA를 세포라인 Molt 4(Human Genetic Mutant Cell Repository사로부터 입수, GM 2219c)로부터 하기 문헌[Stetler et al., Proc, Nat Acad. sci. (1982), 79 : 5966-5970]에 기술된 기법을 사용하여 추출한다.
b. 클론된 글로빈 유전자의 제조
정상적인 β-글로빈 유전자의 1.9kb BamH-1 단편을 코스미드(cosmid)pFC11로부터 단리시켜 pBR328의 BamH I부위에 삽입시킨다. Soberon et al., Gene(1980) 9 : 287-305. 이 단편은 합성 40-머탐침으로 하이브리드화 시킬 부위를 에워사고 있는데, 첫 번째 및 두 번째 액손(exon), 첫 번째 인트론(intron) 및 유전자의 5'측면 서열을 포함하고 있다[Lawn et al., Cell (1978(, 15 : 1157-1174]. 이 클론은 pBR 328 : HbA로 표기하며 ATCC 제39698호로 1984년 5월 25일에 기탁하였다.
상응하는 β-글로빈의 경상 적혈구 대립인자의 1.9kbH I 단편은 코스미드 pFC 12로부터 단리시켜 상기 기술한 바대로 클론한다. 이 클론은 pBR 328 : Hbs로 표기하여 ATCC 제39699호로 1984년 5월 25일에 기탁하였다.
각 재조합 플라스미드를 형질전환시켜 이. 콜라이 MM 294(ATCC 제39,607호)에서 증식시킨다.
C. 클론된 글로빈 유전자를 Mst II로 분해시키는 방법
pBR 328 : HbA 및 pBR 328 : Hbs 각각을 전체 100μ씩으로, 각기 150mM NaCl, 12mM 트리스 HCL(pH 7.5), 12mM ㎎Cl2, 1mM 디티오트레이톨(DTT) 및 100㎍/㎖ 소의 혈청 알부민(BSA)으로 이루어진 용액 200㎕ 중, 37℃에서 16시간동안 Mst II 20단위 (New England Biolabs)로 분해시킨다. 생성물은 각각 pBR 328 : HbA.Mst II 및 pBR 328 : HbS II로 표기한다.
III. 폴리머라제 연쇄 반응
60mM 나트륨 아세테이트, 30mM 트리스 아세테이트 및 10mM 마그네슘 아세테이트로 이루어진 완충액(pH 8.0) 100㎕에, 프라이머 A(서열 CACAGGGCACTAACG)100피코몰, 프라이머 B(서열 d(TTTGCTTCTGACACA)) 100피코몰 및 dATP, dCTP, dGTP 및 TTP 각 1000피코몰를 함유하는 용액 2㎕로 가한다. 상기 기술한 DNA이 하기원중 하나를 가한다.
전 인체의 야생형 DNA 10㎍(반응 I)
pBR 328 : HbA 0.1피코몰(반응 II)
pBR 328 : HbS 0.1피코몰(반응 III)
pBR 328 : HbA/Mst II 0.1피코몰(반응 IV)
pBR 328 : HbS/Mst II 0.1피코몰(반응 V)
표적 DNA부재(반응 VI)
각각의 생성된 용액을 100℃로 4분 동안 가열한 다음 실온으로 2분동안 냉각시키고, 이. 콜라이 DNA 폴리머라제의 클레노우 단편 4단우를 합유하는 1㎕를 가한다. 각 반응을 10분동안 수행한 후, 프라이머, 뉴클레오타이드 및 DNA를 가하여, 가열하고, 냉각시키고, 폴리머라제를 가하는 사이클을, 반응(I)의 경우에는 19회, 반응(II) 내지 반응(VI)의 경우에는 4회 반복 수행한다.
각 반응의 첫 번째 사이클전에 그리고 최종 사이클후에 취한 반응(I) 및 반응(II)의 분취량 4마이크로리터를, 0.089M 트리스-브레이크 완충액(pH 8.3) 및 2.5mM EDTA 중의 12% 폴리아크릴아미드겔에 올려 놓는다. 겔을 25Volts/㎝에서 4시간동안 전기 영동시켜, 고체 상 지지체로 제공된 나일론 막에 옮긴 다음, 30% 포롬아미드, 3×sspE, 5×Denhardt's 5% 나트륨 도데실 설페이트(pH 7.4)중에서, 표준 기법으로 제조한 하기 서열을 갖는 5'-32P로 표지된 40bp 합성 단편으로 탐침한다 : 5'd(TCCTGAGGAGAAGTCTGCCGTTACTGCCCTGTGGGGCAAG) 3'
제2도는 반응(I) 및 반응(II)의 경우 탐침된 나일론 막의 방사선 사진이다. 레인 1은 그의 본쇄 하나가 상기 탐침에 상보적인 58-bp 합성 단편 조절자 0.1피코몰이다. 레인 2는 반응(I)의 첫 번째 증대 사이클을 수행하기 전의 4㎕이다. 레인 3은 반응(I)의 증대 사이클을 20회 수행한 후에 4㎕이다. 레인 4는 반응(II)의 증대 사이클을 5회 수행한 후의 4㎕이다. 레인 5는 알파-32P-dNTPs 및 폴리머라제로 표지한 pBR 322(New England Biolabs) Fok I(New England Biolabs) 분해물로 이루어진 분자량 표준이다. 레인 3은 사이클을 20회 수행한 후에 반응 혼합물(I)이 적절한 분자량이 갖는 특정 서열을 상당량 함유하지만 다른 탐지 가능한 생성물을 함유하지 않음을 보여준다. 레인 4에서 볼 수 있는 바와 같이, 사이클을 5회 수행한 후에 반응 혼합물(II)은 이 생성물 뿐만 아니라, 출발핵산 및 다른 생성물을 함유한다.
반응 사이클로 4회 수행한 후의 반응(II) 내지 반응(VI)의 분취량 5.0㎕에 상기 기술한 각 프라이머 5피코몰을 가한다. 용액을 4분 동안 100℃로 가열한 다음 실온이 되게 한다. 알파-32P-dATP 알파-32P-dCTP 알파-32P-dGTP 및 알파-32P-dTTP 각각 3피코몰 및 클레노우 단편 4단위를 가한다. 최종 용적 10㎕중, 상기 기술한 염 농도에서 반응을 10분 동안 진행시킨다. 폴리머라제 활성은 60℃에서 20분 동안 가열하l면 상실된다. 반응(II) 내지 반응(VI)의 분취량 4㎕를 0.089M트리스-브레이트 완충액 (pH 8.3) 및 2.5mM EDTA 중의 12% 폴리아크릴아미드 결상에 올려 놓는다. 겔을 25Volts/㎝에서 4시간 동안 전기 영동시킨 후 사진을 찍는다.
제3도는 전기 영동사진이다. 레인 1은 분자량 표준이고, 레인 2는 반응(II이며, 레인 3은 반응(III)이고, 레인 4는 반응(IV)이며 레인 5는 반응(V)이다. 조절자로 DNA를 갖지 않은 반응(VI)에 대한 레인은 어느 레인에서도 그 상(image)이 나타나지 않는다. 도면으로부터, 표적 DNA로부터 예상된 94-bp 단편은 손상되지 않은 β-글로빈 DNA 서열이 증대 가능한 경우, 즉 pBR 328 : HbA(레인 2), pBR 328 : HbS(레인 3) 및 pBR 328 : HbS/Mst II(레인 5)에서만 나타남을 알 수 있다. Mst II 분해로 pBR 328 : HbAd[사 94-머 서열이 증대될 수 없도록 절단되어, 94-머 서열은 레인 4에서 나타나지 않았다. 대조적으로, pBR 328 : HbS에서 94-머 서열은, 플라스미드를 Mst II로 분해시키는 경우, 절단되지 않으므로 레인 5에서 볼 수 있는 바대로 증대 가능하다.
제4도는 94-bp 서열을 증대시킴에 있어 3개의 사이클의 연쇄 반응을 설명한 것이다. PCO1 및 PCO2는 프라이머 A 및 B이다. 오른편에 있는 숫자는 사이클을 나타내고, 왼편에 있는 숫자는 특정 분자가 생성되는 사이클 번호를 나타낸다.
[실시예 3]
이 실시에는 인체의 헤모글로빈 유전자에 있어 대립인자의 Mst II 부위를 함유하는 110bp 서열의 증대를 설명하는 것이다.
실시예 2의 기법으로 재조한 전인체의 DNA 1.0마이크로그램, 프라이머 d(ACACAACTGTGTTCACTAGC) 100피코몰 및 프라이머 d(CAACTTCATCCACGTTCACC)100피코몰 전체를 하기 성분으로 이루어진 용액 100㎕에 용해시킨다.
각 1.5mM인 4개의 데옥시 리보뉴클레오사이드 트리포스페이트.
30mM 트리스 아세테이트 완충액(pH 7.9)
60mM 나트륨 아세테이트
10mM 마그네슘 아세테이트
0.25mM 디티오트레이톨
용액을 1분 동안 100℃로 가열한 다음 1분 동안 재빨리 25℃로 되게한 후, DNA 폴리머라제의 클레노우 단편 2.5단위를 가한다. 폴리머라제 반응을 25℃에서 2분 동안 진행시킨 후, 가열하여, 냉각시키고, 클레노우를 가하는 사이클 반응을 목적으로 만큼 자주 반복 수행한다.
각 사이클을 70% 효율로, 15회 반복 수행하면 β-글로빈 유전자의 목적하는 110bp 단편 1.4펨토몰이 합성된다.
[실시예 4]
이 실시예는 인체의 헤모글로빈 유전자에 있어 대립인자의 Mst II 부위를 함유하는 240pb 서열의 증대를 설명하는 것이다. 이 서열은 Nco I, Hinf I 및 Mst II 제한 부위들을 함유한다.
pBR 328 : HbA 0.1 피코몰을 함유하는 60mM 나트륨 아세테이트, 30mM 트리스 아세테이트 및 10mM 마그네슘 아세테이트의 혼합물(pH 8.0) 100㎕에 하기 성분을 함유하는 용액 A 2㎕를 가한다 :
d(GGTTGGCCAATCTACTCCCAGG) 프라이머 100피코몰
d(TAACCTTGATACCAACCTGCCC) 프라이머 100피코몰
dATP, dCTP, dGTP 및 TTP 각각 1000피코몰.
실시예 2에 기술된 기법으로 2개의 프라이머를 제조한다. 용액을 4분동안 100℃로 가열한 다음 2분 동안 주변 공기중에서 냉각시킨 후, 이. 콜라이 DNA 폴리머라제의 클레노오 단편 4 단위를 함유하는 1㎕를 가한다. 반응을 10분동안 진행시킨 후, 용액 A를 가하고, 가열하여 냉각시키고, 폴리머라제를 가하는 사이클 반응을 3회 반복수행한다. 반응물의 분취량 5.0㎕에 상기 기술한 올리고뉴클레오타이드 프라이머 각 5피코몰을 가한다. 용액을 4분 동안 100℃로 가열한 다음 주변 온도로 되게한 후, 알파-32P로 표지된 데옥시리보뉴클레오사이드 트리포스페이트각 3피코몰 및 플레노우 단편 4 단위를 가한다. 최종 용적 10㎕중, 상기 기술한 염 농도에서 반응을 10분동안 진행시킨다. 폴리미라제 활성은 60℃에서 20분 동안 가열하면 상실된다. 분취량 2㎕를 Nco I, Mst II 또는 Hinf I로 분해시킨 후 0.089M 트리스-보레이트 완충액(pH. 8.3) 및 2.5mM EDTA중의 중의 12% 폴리아크릴아미드 겔상에 올려놓는다. 겔은 25Volts/㎝에서 4시간 동안 전기 영동시킨 후 사진을 찍는다. 제5도는 전기 영동 사진인데, 여기에서 레인 1은 분자량 표준이고, 레인 2는 효소로 분해시키지 않은 것(상되지 않은 240bp)이며, 레인 3은 Nco I으로 분해시킨 것(131 및 109bp)이고, 레인 4는 Mst II로 분해시킨 것(149 및 91bp)이며, 레인 5는 Hinf I로 분해시킨 (144 및 96bp)이다. 사진은 240bp 서열의 증대와 일치한다.
[실시예 5]
이 실시예는 겸상절혁구 빈혈증을 연쇄적 분쇄에 의해 탐지하고자 하는 본 발명의 방법의 용도를 설명하는 것이다.
[올리고데옥시리보 뉴클레오타이드의 합성 및 탈인산화]
하기 서열을 갖는, 표지한 DNA 탐침 RS06 :
5'*CTGACTCCTGAGGAGAAGTCTGCCGTTACTGCCCTGTGGG 3'
(여기에서, *는 표지를 나타냄), 및 하기 서열을 갖는 표지하지 않은 블록 올리고머 RS10 :
3'GACAGAGGTACCTCTCAGACGGCAATGACGGGACACCC 5'
(RS06과는 3개의 염기쌍이 잘못 짝지어져 있다)을 실시예 2(Ⅰ)에 기술된 방법에 따라 합성한다. 탐침 RS06은, 이 RS06 5피코몰을, 70mM 트리스완충액 (pH 7.6), 10mM ㎎Cl21.5mM 스페르민 및 2.5mM 디티오트레이톨을 함유하는 반응물 용적 40㎕중, 37℃에서 90분 동안 T4 폴리뉴클레오타이드 키나제(New England Biolabs) 4단위 및 γ-32P-ATP(New England Nuclear, dir 7200Ci/mmole) 50피코올과 접촉시켜 표지한다. 전체 용적을 25mM EDTA로 100㎕로 조정한 다음, 트리스-EDTA(TE) 완충액(10mM 트리스 완충액, 0.1mM EDTA pH 8.0)으로 채운 바이오래드(BioRad)사 제품인 Bio Gel P-4스핀 투석 칼럼상에서 하기 문헌[Maniatis et al,(1982), 464-465]에 기술된 방법에 따라 정제한다. 표지된 탐침은 트리스-붕산-EDTA(TBE)완충액(89mM 트리스, 89mM 붕산, 2.5mM EDTA pH8.3)중의 18% 폴리아크릴아미드 겔(19 : 1아크릴아미드 : BIS, BioRad사)사에서 500vhr 동안 전기 영동시켜 더 정제한다.
사진을 찍어 위치를 측정한 후, 표지된 탐침을 함유하는 겔의 부위를 절단하여, 분쇄하고, TE 완충액 0.2㎖내로 4℃에서 밤새 용출시킨다. 반응 새성물을 TCA로 침전시킨 결과 고유 활성은 4.9 Ci/mmole이고 최종 농도는 20pmole/㎖이다.
표지하지 않은 RS10 블록 오리고머는 200pmole/㎖의 농도로 사용된다.
[세로 라인으로부터 인체 게놈의 DNA의 단리]
고분자량을 갖는 게놈의 DNA를, 본질적으로 하기 문헌[Stetler et al.,(1982), 5966-5970(Molt 4의 경우) 및 Maniatis et al.,(1982), 280-281]에 기술된 방법을 사용하여 임파 세포라인 Molt 4, SC-1 및 GM2064로부터 단리시킨다.
Molt 4(Human Mutant Cell Repository GM 2214C)는 정상적인 β-글로빈에 동형 접합성인 T세포라인이고, SC-1(1985년 3월 19일 ATCC에 기탁)은 겸상적혈구 빈혈중에 동형접합성인, EBV로 형질전환된 B 세포라인이다. GM 2064(Human Mutant Cell Repository GM 2064)는 원래 태아 헤모글로빈의 유전지속체(HPFH)에 각각 동형접합성인 것으로부터 단리시킨 것이며 베타-또는 델타-글로빈 유전자 서열은 함유하지 않는다. 모든 세포라인은 10% 송아지 태아혈정을 사용하여 RPMI-1640내에 저장한다.
[임상 혈액 샘플로부터 인체 게놈의 DNA의 단리]
알려진 겸상적혈구 담체(As)로부터 CH12로 표기된 임상 혈액 샘플을 미합중국 캘리포니아 오클랜드 소재의 Children's Hospital에 근무하는 Bertram Lubin 박사로부터 입수하였다. 게놈의 DNA는, 주로 말초혈액 임파구 세포(peripheral blood lymphocytes)로 이루어진 연층 분획(buffy coat fraction)으로부터, 하기 문헌[Nunberg et al.,, 5553-5556(1978)에 기술된 방법의 수정방법을 사용하여 제조한다.
세포를 트리스-EDTA-NacL(TEN) 완충액(10mM 트리스 완충액 pH 8, 1mM EDTA 10mM NaCl) 5㎖에 재현탁시켜, 0.2㎎/㎖ 프로테이나제 K, 0.5% SDS로 조정한 다음 37℃에서 밤새 배양한다. 과염소산 나트륨을 0.7M로 가한 후 용균물(lysate)을 실온에서 1 내지 2시간 동안 부드럽게 진탕시킨다. 용균물을 페놀/클로로포롬(1 : 1) 30㎖로 추출한 후, 클로로포름 30㎖로 추출하고, 이어서 핵산을 에탄올 침전시킨다. 펠렛을 TE 완충액 2㎖에 재현탁시켜 RNase A를 0.005㎎/㎖로 가한다. 37℃에서 1시간 동안 분해시킨 후, DNA를 페놀, 페놀/클로로포롬 및 클로로포름 각 동용적으로 1회씩 추출한 후, 에탄올 침전시킨다. DNA를 TE 완충액 0.5㎖에 재현탁시켜, 260nm에서의 흡광도로 농도를 측정한다. 선택적으로 β-그로빈 서열을 증대시키기 위한 폴리머라제 연쇄 반응.
게놈의 DNA2 마이크로그램을, 10mM 트리스 완충액(pH 7.5), 50mM NaCl 10mM MgCl2서열의 프라이머 A d(CACAGGGCACTAACG) 150피코몰 및 서열의 프라이머 B d(CTTTGCTTCTGACACA) 150피코몰을 함유하는 초기 반응물 용적 100㎕에서 증대시켜 증발되지 않게 광유(mineral oil) dir 100㎕를 바른다.
각 DNA 샘플은 하기의 3단계로 이루어진 사이클을 15회 반복 수행하여 증대시킨다.
1) 95℃로 고정시킨 열 차단 세트내에서 2분 동안 변성시킨다.
2) 즉시 30℃로 고정시킨 열 차단 세트로 옮겨 2분 동안 보관하여 프라이머 및 게놈의 DNA가 어닐링되게 한다.
3) 10mM 트리스(pH 7.5), 50mM NaCl, 10mM MgCl2및 4mM 디티오트레이톨로 이루어진 완충액중에서, 이.콜라이 DNA 폴리머라제 I의 클레노우 단편(New England Biolabs) 5단위, dATP, dCTP, dGTP 및 TTP 각 1나노몰을 함유하는 용액 2㎕를 가한다. 이 증대반응을 30℃에서 10분 동안 진행시킨다.
최종적으로 사이클을 반복수행한 후, 반응은, 95℃에서 2분동안 가열하여종결시킨다. 광유를 클로로포름 0.2㎖로 추출하여 폐기한다. 최종의 반응물 용적은 130㎕이다. 증대된 게놈의 dna를 탐침 및 Dde I/Hinf I를 사용하여 하이브리드화/분해시키는 반응.
증대된 게놈의 DNA 45마이크로리터를 에탄올 침전시켜 TE 완충액 동용적에 재현탁시킨다. 10마이크로리터(게롬 DNA 154ng의 증대 전 등가물 함유)를 1.5㎖ 미크로퓨지 튜브에 분해하여 TE 완충액 20㎕로 최종 용적이 30㎖가 되게 한다. 샘플에 광유를 바른 후 95℃에서 10분 동안 변성시킨다. 표지된 RS06탐침 0.02피코몰을 함유하는 0.6M NaCl 10마이크로리터를 튜브에 가하여 부드럽게 혼합하고, 즉시 56℃로 고정시킨 열 차단 세트에 옮겨 1시간동안 보관한다. 표지하지 않은 RS10 블록 오리고머(0.8피코몰) 4마이크로 리터를 가하여, 동일한 온도에서 10분동안 더 하이브리드화를 계속한다. 60mM MgCl2/0.1% BSA 5마이크로리터 및I 1㎕(10단위, New England Biolabs)를 가하여, 제어닐링된 DNa를 56℃에서 30분동안 분해시킨다.I 1마이크로리터(10단위, New England Biolabs)를 가하여, 30분동안 더 배양한다. 반응은 75mM EDTA 4㎕ 및 추적 염료 6㎕를 최종 용적 61㎕가 되게 가하면 중지된다.
광유를 클로로포롬 0.2㎖로 추출하고, 반응 혼합물(45ng 게놈 DBA) 18㎕를 Hoeffer SE 200 장치내에서 30% 폴리아클리아미드 미니-겔(19 : 1, BioRad) 상에 올려놓는다. 겔을 약 300Volts에서 1시간동안, 브로모페놀블루 염료가 원점에서 3.0㎝ 이동할때가지, 전기 영동시킨다. 겔의 상부 1.5㎝를 제거하고 남아있는 겔을 -70℃에서 4일간 1개의 보력 스크린 (intensification screen)을 사용하여 노출시킨다.
[사진 설명(제9도)]
각 레인은 증대된 게놈 DNA 45ng을 함유한다. 레인 A는 Molt 4 DNA를 함유하고 : 레인 B는 CH12를 함유하여 : 레인 C는 SC-1을 함유하고 : 레인 D는 GM 2064를 함유한다. Molt 4는 세포당 βA유전자 2복 제본을 갖는 정상적인 개개의 유전자형(AA)을 나타내고, CH12는 세포당 하나의 βA및 하나의 βS유전자를 갖는 겸상적혈구 세포 담체로부터의 임상 샘플(AS)을 나타내며, SC-1은 세포당 βS유전자 2복제본을 갖는 겸상적혈구 세포 개개의 유전자형(SS)을 나타낸다. GM2064는 베타 - 또는 델타-글로빈 서열을 함유하지 않는데, 음성 조절자(negative control)로 존재한다.
사진에서 볼 수 있는 바와같이,I로 절단한 βA-특이성 옥타머는 βA-유전자를 함유하는 그러한 DNS's에서만 존재(레인 A 및 B)하고I로 절단한 βS-특이성 트리머는 βS유전자를 함유하는 그러한 DNA's에서만 존재(레인 B 및 C)한다. 트리머 및 옥타머 둘 모두의 존재(레인 B)는 겸상적혈구 세포담체에 대한 특징이고 옥타머만을 갖는 정상적인 개개(레인 A)와 구별 가능하고 트리머만을 갖는 개개에 의해 영향을 받은 겸상 적혈구 세포(레인 C)와도 구별 가능하다.
비교용으로, 증대되지 않은 게놈 DNA를 사용하여 상기 기술된 실험을 반복수행하면 증대가 탐지 민감도를 적어도 1000배까지 증가시킴을 알 수 있다.
[실시예 6]
이 실시예는 표지된 탐침을 필요치 않고서 겔상에서 전인체의 DNA에서의 전체적으로 정제되지 않은 단일 복제 유전자의 직접적인 탐지법을 설명하는 것이다.
실시예 3에 기술된 기법을 사용하며, 베타-글로빈 유전자의 첫 번째 액손에서의 서열로부터 110-bp 단편을, 사이클로 20회 반복수행한 후 전 인체의 DNA 10㎍으로부터 증대시킨다. 사이클을 20회 반복수행한 후 생성된 이 110-bp 단편을 에티디움 브로마이드로 염색을 겔상에서 쉽게 가시화된다.
[실시예 7]
A. 인체의 베타-글로빈 A 빈혈증으로부터 1.9Kb 삽입물, 알파-32p-dNTP(500c : mole) 각 50 나노몰 및 실시예 3에서 사용된 프라이머 각 1나노몰을 함유하는 pBR328 100펨토몰 전체를 30mM 트리스-아세테이트(pH 7.9), 60mM 나트륨 아세테이트, 100mM 디티오트레이톨 및 10mM 마그네슘 아세테이트를 함유하는 용액 100㎕에 용해시킨다. 이 용액을 2분동안 100℃로 되게 한 다음 1분동안 25℃로 냉각시킨다.DNA 폴리머라제 1의 클레노아 단편 4.5 단위 및 무기 피로포스파타제 0.09 단위를 함유하는 1㎕ 전체를 가하여 반응 혼합물에서 피로포스페이트의 축적(build-up)가능성을 방지하고, 반응을 25℃에서 2분동안 진행시킨 후, 가열하여, 냉각시키고, 효소를 가하는 사이클반응을 9회 반복수행한다. 분취량 10㎕를 취하여 600mM EDTA 1㎕에 가한 후 각 합성사이클을 수행한다. 각각은 90mM 트리스-보레이트 및 2.5mM EDTA중의 14% 폴리아크릴아미드겔상 pH 8.3 및 24 Volts/㎝에서 2.5시간 동안 분석한다. 완성된 겔을 0.5㎍/㎖ 에티디움 브로마이드를 가하면서 동일한 완충액에 20분동안 담근후, 원래의 완충액으로 세척하고, 적색 필터를 사용하여 UV광에서 사진을 찍는다.
생성된 110-bp 단편을 UV 광 하에서 겔로부터 절단하여 세렌코브 방사(cerenkov radiation)에 의해 카운트된32p를 혼입시킨다. 하기 방정식 : pmoles/10㎕=0.01[(1+y)N-yN-1] {여기에서, N은 사이클의 수이고 y는 사이클당 분획 수율이다}의 데이터를 적절히 하기 위한 시도로 y=0.619일 경우 최적이다. 이것으로 상당한 양의 증대가 발생했음을 알 수 있다.
B. 각 dNTP 100 나노몰을 반응물 100㎕에 가하고, 방사선 표지를 사용하지 않으며, 분취량을 각 사이클에서 취하지 않는 것을 제외하고는 상기 실험을 반복 수행한다. 사이클을 10회 반복 수행한 후 2분동안 비등시켜 반응을 종결시키고 57℃에서 1시간동안 재하이브리드화를 수행한다. 110-bp 생성물의 서열은, 분위량 8㎕에 소혈청 알부민(25㎎/㎖) 1㎕ 및 적절한 제한 효소()1㎕를 가한 후 37℃에서 15시간동안 반응시킴에 의한 제한 분석으로 확인한다. PAGE도 상기 기술한 바와 같이 수행한다.
[실시예 8]
이 실시예는 pBR328 및 pBR322의 여러 단편을 증대시키기 위한 상이한 프라이머의 용도를 설명한 것이다.
A. 하기 프라이머 : d(TTTGCTTCTGACACAACTGTGTTCACTAGC) 및 d(GCCTCACCACCAACTTCATCCACGTTCACC)를 사용하는 것을 제외하고는, 실시예 7A에 기술된 실험을 반복수행하여 pBR328의 130-pbeks단편을 수득한다.
B. 하기 프라이머 : d(GGTTGGCCAATCTACTCCCAGG) 및 d(TGGTCTCCTTAAACCTGTCTTG)를 사용하는 것을 제외하고는, 실시예 7A에 기술된 실험을 반복수행하여 pBR328의 262-bp 단편을 수득한다. 반응시간은 사이클당 20분이다.
C. 인체의 베타-글로빈 S 대립인자로부터의 1.9Kb 삽입물을 함유하는 pBR328의II 분해물 100 펨토몰을 최초의 주형으로 사용하는 것을 제외하고는, 실시예 8B에 기술된 실험을 반복 수행한다. 이 플라스미드를, 증대시킬 서열 내부는 제외하고는,II로 수회 절단한다. 또한 하기 프라이머 : d(GGTTGGCCAATCTACTCCCAGG) 및 d(TAACCTTGATACCAACCTGCCC)를 사용하여 240-bp단편을 수득한다.
D. pBRr322의 Nru I 분해물 100 펨토몰을 주형으로 사용하고, 각 dNTP 200 나노몰을 반응물 100㎕에 사용하고, 하기 프라이머 : d(TAGGCGTATCACGAGGCCCT) 및 D(CTTCCCCATCGGTGATGTCG)를 사용하는 것을 제외하고는 실시예 7에 기술된 실험을 반복 수행하여 pBR322로부터 500-pb 단편을 수득한다. 반응 시간은 37℃에서 사이클당 20분이다. 최종의 재하이브리드화는 57℃에서 15시간동안 수행한다. 전기영동은 4% 아가로오스 겔 상에서 행한다.
[실시예 9]
이 실시예는 생체의 돌연변이가 증대된 도입되는 본 발명의 방법을 설명하는 것이다.
A. 40mM 트리스(pH 8), 20mM MgCl2, 5mM 디티오트레이틀 및 5㎎/㎖ 소혈청 알부민으로 이루어 전 용액 100㎕중에서, Nru I으로 선형화한 pBr322 100 펨토몰, 75-BP 단편을 생성시킬 하기 프라이머 각 1나노몰 : d(CGCATTAAAGCTTATCGATG) 및 d(TAGGCGTATCACGAGGCCCT) 및 각 dNTP 100 나노몰을 합한다. 혼합물을 1분동안 100℃로 되게 한다음, 23℃의 수욕에서 0.5분동안 냉각시키고, 클레노우단편 4.5 단위 및 무기 피로포스파타제 0.09단위를 가한 후, 반응율 3분동안 진행 시킨다. 가열하여, 냉각시키고, 효소를 가하는 사이클반응을 9회 반복 수행한다. 동결시킴에 의해 10회째 반응 사이클을 종결시킨 후, 반응혼합물 분취량 8㎕를 에티디움 브로마이드로 가시화한 4% 아가로스 겔에 올려 놓는다.
b. 하기 올리고뉴클레오타이드 프라이머: d(CGCATTAAAGCTTATCGATG) 및 d(AATTAATACGACTCACTATAGGGAGATAGGCGTATCACGAGGCCCT)를 사용하는 것을 제외하고는, 실시예 9A에 기술된 실험을 반복 수행한다. 이들 프라이머는 101-bp 단편을 생성시키도록 계획되어 있으며, 두 번째에 기술한 프라이머의 26뉴클레오타이드는 pBR322에는 존재하지 않는다. 이들 뉴클레오타이드는 T7 프로모터의 서열을 나타내는데, 20개의 상보적인 염기 및 26-염기 5' 증대를 갖는 프라이머를 사용하여 pBR322로부터의 75-bp 서열에 부착시킨다. 이 절차를 수행하는 데에는 2시간 미만이 소요되며 pBR322 100펨토몰로부터 비교적 순수한 102-bp 단편 2 피코몰이 생성된다.
T7 프로모터는 RNA 전사를 일으키는데 사용될 수 있다. T7 폴리머라제를 101-bp 단편에 가하여 일본쇄 RNA를 제조할 수도 있다.
C. 하기 올리고뉴클레오타이드 프라이머: d(TAGGCGTATCACGAGGCCCT) 및 d(CCAGCAAGACGTAGCCCAGC)를 사용하는 것을 제외하고는, 실시예 8D에 기술된 실험을 반복 수행하여 pBR322로부터 1000-BP 단편을 수득한다.
d. 하기 올리고뉴클레오타이드 프라이머: D(TAGGCGTATCACGAGGCCCT) 및 d(AATTAATACGACTCACTATAGGAGATAGGCGTATCACGAGGCCCT)를 1026-bp 단편을 제조하기 위해 사용 {여기에서, 두 번째 기술한 프라이머의 26뉴클레오타이드는 pBR322에 존재하지 않으며 상기 기술한 T7 프로모터를 나타낸다}하는 것을 제외하고는, 실시예 9C에 기술된 실험을 반복수행한다. 프로모터는 pBR322로부터 1000-bp 단편에 인접하게 삽입되어 있다.
결과로, 프라이머가 주형 서열에 완전하게 짝을 이루지 않지만, 그럼에도 불구하고 충분히 하이브리드화하여 효소적으로 증대시켜, 상응하는 원 주형의 서열보다 프라이머의 서열을 함유하는 장쇄 생성물을 수득할 수 있음을 알 수 있다. 장쇄 생성물은 두 번째의 프라이머를 위한 주형으로 제공되어 생체내 돌연변이를 유도한다. 추가의 사이클에서 이 돌연변이는, 더 이상의 짝을 못이룬 프라이머는 필요로 하지 않기 때문에, 효율의 감소 없이 증대된다. 이 경우, 그의 5' 말단상에서 비-상보적인 증대를 수반하는프라이머는 새로운 서열을 복제될 주형 서열에 인접한 생성물에 삽입시키는데 사용된다.
E. 폴리머라제의 반응은 피로포스페이트를 생성하며 이론상 가역적이기 때문에 (Komberg, A., DNA Replication, W, H. Freeman, San Francisco, 1980, 잠재적 피로포스포르분해를 방지하기 위해 함유한 무기 피로포스페이트의 효과를 측정하였다. 반응을 더하기 및 빼기 피로포스파타제의 질적 롤리아크릴아미이드겔 전기영동 시험은 이 효소의 함유결과 생성물의 동종서에 작기는 하나 확실한 증가를 입증하였다.
[실시예 10]
이 실시예는 단일 복제 유전자의 증대시 배경을 감소시키기 위해 사용하는 프라이머의 네스트 세트(nested set)를 설명하는 것이다.
야생형 β-글로빈 대립인자에 동형 접합성인 전 인체 DNA를 하기와 같은 증대 사이클을 20회 반복 수행한다 : 300mM 트리스-아세테이트(pH 7.9), 60mM 나트륨 아세테이트, 10mM 디티오트레이톨 및 10mM 마그네슘 아세테이트로 이루어진 용액 100㎕중의, DNA 10㎍ 하기 프라이머 d(ACACAACTGTGTTCACTAGC) 및 d(CAACTTCATCCACGTTCACC) 각 200 피코몰 및 각 dNTP 100나노몰 전체를 100℃로 1분동안 가열한 다음, 25℃로 1분동안 냉각시키고, 2분동안 클레노우 단편 2 단위로 처리한다. 가열하고, 냉각하여 클레노우 단편을 가하는 사이클 반응을 19회 반복 수행한다. 반응 혼합물로부터 분취량 10㎕를 취하여 하기 프라이머:D(CAGACACCATGGTGCACCTGACTCCTG) 및d(CCCCACAGGGCAGTAACGGCAGACTTCTCC) 각각을 사용하여 증대 사이클을 10회 더 반복수행하면, 상기 생성된 110-bp 단편내에 함유되어 있는 58-bp 단편이 증대된다. 이 최종적인 증대 사이클 10회는, 분취량 10㎕를 가가 dNTP 100 나노몰 및 각 프라이머 200 피코몰을 함유하는 상기 기술한 신선한 트리스-아세테이트 완충액 90㎖에 희석시켜 수행한다. 반응 조건은 상기 기술한 바와 같다. 사이클을 10회 수행한 후, 분취량 10㎕(원 DNA 100 나노그람에 사용)를 6% Nusieve(FMC Corp)아가로수겔 상에 올려놓고 에티디움 브로마이드를 사용하여 가시화한다.
제10도는 UV 광을 조사한 후 본 분야에서 알려진 바대로 적색 필터를 통해 사진을 찍은 겔을 설명한 것이다. 레인 1은 분자량 표지물이다. 레인 2는 상기 기술한 반응의 분취량이다. 레인 3은 원래의 야생형 DNA를 증대시키기 전에I으로 절단한 것을 제외하고는, 상기 기술한 것과 동일한 반응의 분취량이다. 레인 4는 겸상 적혈구 베타글로빈 대립인자에 동형 접합성인 인체 DNA를 증대시키기 전에I으로 처리(감상 적혈구 대립인자는 여기에서 증대될 단편에서 Dde I 부위를 함유하지 않는다)한 것을 제외하고는, 상기 기술한 것과 동일한 반응의 분취량이다. 레인 5는 연어 정액 DNA를 인체 DNA 대신 사용한 것을 제외하고는, 상기 기술한 것과 동일한 반응의 분취량이다. 레인 6은 분취량을 Dde I으로 처리한 후 증대(I은 58-bp 야생형 27-bp 및 31-bp 단편으로 전환시킨다) 시킨 것을 제외하고는, 상기 기술한 것과 동일한 반응의 분취량이다. 레인 7은 레인 4 물질을I으로 처리한 후 증대(58-bp 겸상적혈구 생성무른I 부위를 함유하지 않는다)시킨 분취량이다.
인체 DNA 1㎍으로부터 단일 복제 유전자의 대표적인 58-bp 단편을 단지 에티디움 브로마이드로 염색한 아가로오스 겔만을 사용하여 탐지하려면 약 500,000배의 증대가 필요하다. 이것은 본 명세서에 기술한 올리고뉴클레오타이드 프라이머의 2개의 네스트 세트를 사용하여 수행한다. 첫 번째 세트는 110-bp 단편을 증대시키고 내부 네스트 세트는 이 생성물의 아-단편을 제10도에서 볼 수 있는, 편리하게 탐지할 수 있는 양까지 증대시킨다. 상기 증대 방법으로 증대될 서열내에 함유되어 있고 다른 프라이머의 증대 생성물에 함유되어 있는 더 짧은 서열을 프라이머를 사용하여 증대시키는 이 방법으로, 하기 문헌[예 : Conner et al.,: 278(1983) 및 Leary et al., PNAS(USA): 4045(1983)]에 기술되어 있는 방사성 동위원소 또는 비-방사성 동위원소 탐침 하이브리드화 방법의 도움없이 베타-글로빈 좌(locus)에서 야생형을 겸상 적혈구 대립인자와 구별할 수 없다.
[실시예 11]
본 발명의 방법은, 환자의 DNA 샘플에서, 예를 들어 클라미디아(chlamydia)와 같은 감염질병에 관련된 특정 서열을 목적하는 증대된 서열을 함유하는 비오티닐화된 하이브리드화 탐침을 사용하여 그리고 상기 미합중국 특허 제4,358,535호에 기술되어 있는 방법을 사용하여 탐지하는데 유용하리라고 기대된다. 비오티닐화된 하이브리드화 탐침은 하기 일반식의 공간 팔(spacer arm)을 통해 비오틴에 결합되어 있는 4'-메틸렌치환 4,5'-8-트리메틸프-소라렌을 갖는 부분적으로 이본쇄인 DNA를 삽입하여 조사(irradation)하여 제조한다.
여기에서 R은 -H 또는 -CHO 그룹이고, R"는 -H이며, X는 1 내지 4의 수이고, y는 2 내지 4의 수이다(EP 156, 287에 기술). 탐침상의 비오티닐 글부의 탐지는 Enzo Bio Chem iNC. 사로부터 상업적으로 수득가능한 스트랩타비딘-산 포스파타제 복합물을 사용하고 제조자에 의해 그의 팜플렛에 제한한 탐지절차를 사용하여 수행할 수 있다. 하이브리드화된 탐침은 탐지 복합물의 결합 및 산 포스파타제에 의해 촉매되는 후속반응(이 반응으로 침전 가능한 염료가 생성된다)에 기인하여 침전된 염색 반점으로 볼 수 있다.
세포라인 SC-1(CTCC #0082)은, ATCC 기탁번호 CRL #87576하에 미합중국 메릴랜드 20852 록크빌 파크로온 드라이브 12301 소재의 아메리칸 타입 컬춰 콜렉션(American Type culture Collection : ATCC)에 1985년 3월 19일자로 기탁하였다. SC-1의 기탁은 ATCC와 이 특허출원의 서명자인 세투스 코포레이션사와의 게약에 의한 것이다.
ATCC와의 약정은 기탁을 기술하고 입증하는 미합중국 특허의 공표시, 또는 미합중국 또는 외국 특허원의 공개시 어느 것이 먼저든 이 세포라인의 후속을 영구히 공개하며, 35 CFR §122 및 2에 의한 청장 규정에 따라(특히 886 OG 638에 관한 37 CFR § 1.14포함) 미합중국 특허 및 상표청장에 의해 권한을 받은 자에게 이 세포라인의 후손을 분양하는 것이다. 본 출원의 서명자는 만약 기탁된 세포주가 적당한 조건하에 배양시 죽거나 분실 또는 파괴할 경우, 즉시 동일 세포주의 생존 배양물로 공식대치할 것이다.
개략적으로, 본 발명은 프라이머 증대 생성물을 제조하고, 이것은 후속적으로 또다른 프라이머 증대 반응시 주형으로 작용할 수 있는 연쇄 반응을 이용하여 1개 또는 그 이상 특정 핵산 서열을 증대시키는 방법을 제공하고 있다. 본 방법은 초기에 아주 소량으로 존재하는 핵산 서열을 검출하는데 특히 유용하다.
Claims (10)
- (a) 하나의 프라이머로부터 합성된 증대 생성물이 이의 상보서열로부터 분리될 경우 다른 프라이머의 증대 생성물을 합성하기 위한 주형으로 작용할 수 있을 정도로 각각 특징 서열의 상이한 본쇄에 상당히 상보적인 것으로 선택된, 각각의 상이한 특정 서열의 증대를 위한 두 개의 올리고뉴클레오타이드 프라이머를, 각각의 핵산 본쇄에 상보적인 각 프라이머의 증대 생성물이 각각의 상이한 서열의 증대를 위해 합성될 수 있는 조건하에서, 본쇄에 처리하고 ; (b) 주형상에서 합성된 프라이머 증대 생성물을 주형으로부터 분리하여 일본쇄 분자를 제조하며 ; (c) 단계(b)에서 제조된 각각의 일본쇄 분자를 주형으로 사용하여 프라이머 증대 생성물을 합성하는 조건하에서, 단계(b)에서 제조된 일본쇄 분자를 단계(a)의프라이머로 처리함을 특징으로 하여, 길이가 동일하거나 상이한 두 개의 분리된 상보 본쇄로 이루어진 핵산 또는 핵산들의 혼합물내에 포함된 하나이상의 특정 핵산 서열을 증대시키는 방법.
- 제1항에 있어서, 단계(b) 및 (c)를 일회이상 반복하는 방법.
- 제1항 또는 2항에 있어서, 단계(b)를 변성방법 또는 효소 헬리카제(helicase)를 사용하여 수행하고, 단계 (a) 및 (c)를 중합반응 유도제 및 4개의 상이한 뉴클레오타이드를 사용하여 수행하는 방법.
- 제1항에 있어서, 단계(a) 및 (c)를 이. 콜라이 DNA 폴리머라제 I, 이. 콜라이 DNA 폴리머라제 I의 클레노우 단편(Klenow fragment), T4DNA 폴리머라제, 내열성 효소 및 역전사 효소중에서 선택된 중합반응 유도제를 사용하여 수행하는 방법.
- 제1항에 있어서, 핵산이 DNA이고 프라이머가 올리고데옥시리보뉴클레오타이드인 방법.
- 제1항에 있어서, 단계(a)에서 사용된 핵산들의 혼합물이 단계(c)에 의해 제조된 선행증대 방법의 생성물인 방법.
- 제7항에 있어서, 사용된 프라이머가 선행증대 방법에서 사용된 프라이머와 상이한 방법.
- 제1항에 있어서, 하나의 프라이머가 증대될 특정 서열에 대해 비상보적인 하나이상의 뉴클레이타오드를 포함하는 방법.
- 제1항에 있어서, 단계 (a) 및 (c)에서 사용된 2개의 프라이머가 적어도 1000 : 1의 프라이머 : 상보본쇄의 몰비로 각각 존재하는 방법.
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US716975 | 1985-03-28 | ||
US791,308 | 1985-10-25 | ||
US06/791,308 US4683202A (en) | 1985-03-28 | 1985-10-25 | Process for amplifying nucleic acid sequences |
US791308 | 1997-01-30 |
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KR860007379A KR860007379A (ko) | 1986-10-10 |
KR900008742B1 true KR900008742B1 (ko) | 1990-11-29 |
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KR1019860002333A KR910008641B1 (ko) | 1985-03-28 | 1986-03-28 | 핵산 서열의 증대, 탐지, 및/또는 클로닝 방법 |
KR1019860002332A KR900008742B1 (ko) | 1985-03-28 | 1986-03-28 | 핵산 서열의 증대 방법 |
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EP (5) | EP0201184B2 (ko) |
JP (1) | JP2622327B2 (ko) |
KR (2) | KR910008641B1 (ko) |
AT (2) | ATE165869T1 (ko) |
AU (1) | AU586233B2 (ko) |
CA (1) | CA1237685A (ko) |
DE (5) | DE502588T1 (ko) |
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IE (3) | IE930228L (ko) |
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1985
- 1985-10-25 US US06/791,308 patent/US4683202A/en not_active Expired - Lifetime
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1986
- 1986-03-26 NZ NZ215605A patent/NZ215605A/en unknown
- 1986-03-26 ES ES553464A patent/ES8706822A1/es not_active Expired
- 1986-03-26 DK DK144886A patent/DK171160B1/da not_active IP Right Cessation
- 1986-03-27 EP EP86302299A patent/EP0201184B2/en not_active Expired - Lifetime
- 1986-03-27 DE DE92201226T patent/DE502588T1/de active Pending
- 1986-03-27 CA CA000505375A patent/CA1237685A/en not_active Expired
- 1986-03-27 IE IE930228A patent/IE930228L/xx unknown
- 1986-03-27 EP EP92201226A patent/EP0502588A2/en not_active Withdrawn
- 1986-03-27 IE IE84286A patent/IE62098B1/en not_active IP Right Cessation
- 1986-03-27 DE DE3687287T patent/DE3687287T3/de not_active Expired - Lifetime
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- 1986-03-27 EP EP92201244A patent/EP0505012B2/en not_active Expired - Lifetime
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- 1986-03-27 AT AT86302299T patent/ATE83505T1/de not_active IP Right Cessation
- 1986-03-27 AU AU55322/86A patent/AU586233B2/en not_active Expired
- 1986-03-27 DE DE198686302299T patent/DE201184T1/de active Pending
- 1986-03-27 DE DE199292201244T patent/DE505012T1/de active Pending
- 1986-03-28 KR KR1019860002333A patent/KR910008641B1/ko not_active IP Right Cessation
- 1986-03-28 KR KR1019860002332A patent/KR900008742B1/ko not_active IP Right Cessation
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1992
- 1992-06-12 JP JP4177779A patent/JP2622327B2/ja not_active Expired - Lifetime
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1993
- 1993-10-02 SG SG111193A patent/SG111193G/en unknown
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1994
- 1994-08-18 HK HK84094A patent/HK84094A/xx not_active IP Right Cessation
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1998
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