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PCR](https://www.sigmaaldrich.com#sequencing) - [관련 제품](https://www.sigmaaldrich.com#products) ## [](https://www.sigmaaldrich.com)생어 시퀀싱이란 무엇인가요? "연쇄 종결 방법"이라고도 알려진 생어 시퀀싱은 DNA의 뉴클레오티드 서열을 알아내는 방법입니다. 이 방법은 1977년 두 차례 노벨상 수상자인 프레드릭 생어와 그의 동료들이 개발하여 '생어 서열'이라는 이름이 붙었습니다. DNA의 일반적인 구조를 검토하려면 __그림 2__를 참조하세요. ## [](https://www.sigmaaldrich.com)생어 시퀀싱은 어떻게 작동합니까? 생어 시퀀싱은 수동으로 수행할 수도 있고, 더 일반적으로는 시퀀싱 기계를 통해 자동화된 방식으로 수행할 수도 있습니다(__그림 1__). 각 방법은 아래에 설명된 대로 세 가지 기본 단계를 따릅니다. ![자동화된 생어 시퀀싱.](https://www.sigmaaldrich.com/deepweb/assets/sigmaaldrich/marketing/global/images/technical-documents/protocols/genomics/sequencing/sanger-sequencing_steps_process_diagram/sanger-sequencing_steps_process_diagram.png "자동화된 생어 시퀀싱.") __그림 1.__자동화된 생어 시퀀싱의 세 가지 기본 단계. ## [](https://www.sigmaaldrich.com)Sanger 시퀀싱 단계 Sanger 시퀀싱에는 세 가지 주요 단계가 있습니다. ### 1. 연쇄 종결 PCR을 위한 DNA 서열 관심 있는 DNA 서열은 특수한 유형의 PCR을 연쇄 종결 PCR이라고 합니다. 연쇄 종결 PCR은 표준 PCR과 동일하게 작동하지만, 디데옥시리보뉴클레오티드(ddNTP)라고 하는 변형 뉴클레오티드(dNTP)를 추가한다는 한 가지 큰 차이점이 있습니다. 표준 PCR의 연장 단계에서 DNA 중합효소는 마지막 뉴클레오타이드의 유리 3'-OH 그룹과 다음 뉴클레오타이드의 5'-인산염 사이의 포스포디에스테르 결합 형성을 촉매하여 성장하는 DNA 가닥에 dNTP를 추가합니다(__그림 2__). 연쇄 말단 PCR에서 사용자는 PCR 반응에서 낮은 비율의 연쇄 말단 ddNTP를 일반 dNTP와 혼합합니다. ddNTP에는 포스포디에스테르 결합 형성에 필요한 3'-OH 그룹이 없기 때문에 DNA 중합효소가 ddNTP를 무작위로 결합하면 연장이 멈춥니다. 연쇄 말단 PCR의 결과는 관심 있는 DNA 서열의 수백만에서 수십억 개의 올리고뉴클레오티드 사본으로, 5'-ddNTP에 의해 무작위 길이(n)로 말단됩니다. 수동 시퀀싱에서는 각각 한 가지 유형의 ddNTP(ddATP, ddTTP, ddGTP, ddCTP)만 섞인 네 가지 PCR 반응이 설정됩니다. 자동화된 생어 시퀀싱에서는 모든 ddNTP가 단일 반응에 혼합되며, 4개의 dNTP는 각각 고유한 형광 라벨을 갖습니다. ### 2. 겔 전기영동에 의한 크기 분리 두 번째 단계에서는 겔 전기영동을 통해 사슬 말단 올리고뉴클레오티드를 크기별로 분리합니다. 겔 전기영동에서는 DNA 샘플을 겔 매트릭스의 한쪽 끝에 넣고 전류를 가하면 DNA가 음전하를 띠기 때문에 올리고뉴클레오타이드가 겔의 반대쪽 양극 쪽으로 당겨집니다. 모든 DNA 단편의 질량 단위당 전하가 동일하기 때문에 올리고뉴클레오타이드가 이동하는 속도는 크기에 의해서만 결정됩니다. 조각이 작을수록 젤을 통과할 때 마찰이 적고 더 빨리 움직입니다. 결과적으로 올리고뉴클레오타이드는 가장 작은 것부터 가장 큰 것 순으로 배열되어 젤을 아래에서 위로 읽게 됩니다. 수동 시퀀싱에서는 4개의 PCR 반응 각각에서 나온 올리고뉴클레오타이드가 젤의 4개의 개별 레인에서 실행됩니다. 이를 통해 사용자는 각 ddNTP에 해당하는 올리고뉴클레오티드를 알 수 있습니다. 자동화된 생어 시퀀싱에서는 모든 올리고뉴클레오타이드가 시퀀싱 기계 내에서 단일 캐필러리 겔 전기영동으로 실행됩니다. ### 3. 젤 분석 및 DNA 염기서열 결정 마지막 단계에서는 간단히 젤을 판독하여 입력된 DNA의 염기서열을 결정합니다. DNA 중합효소는 제공된 프라이머에서 시작하여 5' ~ 3' 방향으로만 DNA를 합성하기 때문에 각 말단 ddNTP는 원래 서열의 특정 뉴클레오티드에 해당합니다(예:, 가장 짧은 단편은 5' 끝에서 첫 번째 뉴클레오티드에서 끝나야 하고, 두 번째로 짧은 단편은 5' 끝에서 두 번째 뉴클레오티드에서 끝나야 하는 등) 따라서 젤 밴드를 가장 작은 것부터 큰 것까지 읽으면 원래 DNA 가닥의 5' ~ 3' 서열을 확인할 수 있습니다. 수동 시퀀싱에서는 사용자가 젤의 4개 레인을 한 번에 모두 읽고 아래에서 위로 이동하면서 레인을 사용하여 각 밴드의 말단 ddNTP의 신원을 확인합니다. 예를 들어, 하단 밴드가 ddGTP에 해당하는 컬럼에서 발견되면 가장 작은 PCR 단편이 ddGTP로 종결되고 원래 서열의 5' 끝에서 첫 번째 뉴클레오타이드는 구아닌(G) 염기를 갖습니다. 자동화된 생어 시퀀싱에서는 컴퓨터가 형광을 사용하여 모세관 젤의 각 밴드를 순서대로 읽고 각 말단의 신원을 ddNTP라고 부릅니다. 간단히 말해, 레이저가 각 밴드의 형광 태그를 여기시키고 컴퓨터가 방출되는 빛을 감지하는 것입니다. 4개의 ddNTP는 각각 다른 형광 라벨로 태그되어 있기 때문에 방출되는 빛은 단말기 ddNTP의 신원과 직접 연결될 수 있습니다. 출력은 크로마토그램이라고 하며, 이는 템플릿 DNA의 길이를 따라 각 뉴클레오타이드의 형광 피크를 보여줍니다. ![DNA 구조 도식. DNA는 두 가닥이 서로 감겨 이중 나선을 형성하는 분자입니다. 각 가닥은 데옥시리보뉴클레오티드(dNTP)라는 분자의 문자열로 구성됩니다.](https://www.sigmaaldrich.com/deepweb/assets/sigmaaldrich/marketing/global/images/technical-documents/protocols/genomics/sequencing/sanger-sequencing_dna-structure/sanger-sequencing_dna-structure.png "DNA 구조 도식. DNA는 두 가닥이 서로 감겨 이중 나선을 형성하는 분자입니다. 각 가닥은 데옥시리보뉴클레오티드(dNTP)라는 분자의 문자열로 구성됩니다.") __그림 2.__DNA 구조 도식. DNA는 두 가닥이 서로 감겨 이중 나선을 형성하는 분자입니다. 각 가닥은 데옥시리보뉴클레오티드(dNTP)라는 분자의 문자열로 구성됩니다. 각 dNTP는 인산염기, 당기, 네 가지 질소 염기\[아데닌(A), 티민(T), 구아닌(G) 또는 시토신(C)] 중 하나를 포함합니다. 한 dNTP의 당과 다음 dNTP의 인산염 그룹 사이의 포스포디에스테르 공유 결합에 의해 선형 방식으로 서로 묶여 있으며, 이 반복되는 당-인산염 패턴이 당-인산염 백본을 구성합니다. 두 가닥의 질소 염기는 상보적인 염기 사이의 수소 결합으로 서로 결합하여 이중 가닥 DNA 나선형을 형성합니다. ## [](https://www.sigmaaldrich.com)생어 시퀀싱 결과를 읽는 방법 생어 시퀀싱 결과를 제대로 읽는 것은 두 개의 상보적인 DNA 가닥 중 어떤 것이 관심 대상인지, 어떤 프라이머를 사용할 수 있는지에 따라 달라집니다. 두 가닥의 DNA가 A와 B이고 A 가닥이 관심 가닥이지만 프라이머가 B 가닥에 더 적합한 경우 출력 단편은 A 가닥과 동일합니다. 반면에 A 가닥이 관심 가닥이고 프라이머가 A 가닥에 더 적합한 경우 출력은 B 가닥과 동일합니다. 따라서 출력은 다시 A 가닥으로 변환되어야 합니다. 따라서 관심 서열이 "TACG"이고 프라이머가 해당 가닥에 가장 적합한 경우 출력은 "ATGC"가 되므로 다시 "TACG"로 변환해야 합니다. 그러나 프라이머가 상보 가닥("ATGC")에 더 적합하다면 출력은 올바른 서열인 "TACG"가 됩니다. 요약하면, 시작하기 전에 무엇을 타겟팅하고 어떻게 거기에 도달할지 알아야 합니다! 이 점을 염두에 두고 전자의 예(TACG - & & t; ATGC - & & t; TACG)를 예로 들어보겠습니다. 디옥시뉴클레오타이드 라벨이 T = 노란색, A = 분홍색, C = 진한 파란색, G = 하늘색이면 프라이머-A, 프라이머-AT, 프라이머-ATG 및 프라이머-ATGC라는 짧은 서열이 됩니다. 전기영동으로 단편을 분리하면 레이저가 길이 순서(분홍색, 노란색, 하늘색, 진한 파란색)에 따라 단편을 읽고 크로마토그램을 생성합니다. 컴퓨터가 문자를 변환하므로 최종 염기서열은 올바른 TACG입니다. ## [](https://www.sigmaaldrich.com)Sanger 시퀀싱 대.. PCR Sanger 시퀀싱과 PCR은 유사한 출발 물질을 사용하며 서로 함께 사용할 수 있지만 어느 쪽도 다른 쪽을 대체할 수는 없습니다. PCR은 DNA를 전체적으로 증폭하는 데 사용됩니다. 우연히 다양한 길이의 단편이 생성될 수 있지만(예: DNA 중합효소가 떨어질 수 있음), 목표는 전체 DNA 서열을 복제하는 것입니다. 이를 위해 "성분"은 표적 DNA, 뉴클레오타이드, DNA 프라이머, DNA 중합효소(특히 PCR에 필요한 고온에서 살아남을 수 있는 Taq 중합효소)입니다. 대체로, Sanger 시퀀싱의 목표는 표적 DNA의 전체 길이까지 가능한 모든 길이의 DNA를 생성하는 것입니다. 그렇기 때문에 PCR 시작 물질 외에 이디옥시뉴클레오타이드가 필요합니다. 상어 시퀀싱 프로토콜을 위한 시작 물질을 생성할 때 상어 시퀀싱과 PCR을 함께 사용할 수 있습니다. PCR은 시퀀싱할 DNA의 많은 사본을 만드는 데 사용할 수 있습니다. 작업할 템플릿이 두 개 이상 있으면 Sanger 프로토콜이 더 효율적입니다. 표적 염기서열의 길이가 1,000 뉴클레오타이드이고 템플릿의 사본이 하나만 있는 경우, 태그가 지정된 1,000개의 단편을 생성하는 데 시간이 더 오래 걸립니다. 하지만 템플릿 사본이 여러 개 있는 경우 이론적으로 태그가 지정된 1,000개의 단편을 모두 생성하는 데 걸리는 시간은 더 짧아집니다. ## 관련 제품 예기치 못한 오류가 발생했습니다. 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