JP2622327B2 - 核酸配列の増幅手段 - Google Patents

核酸配列の増幅手段

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Publication number
JP2622327B2
JP2622327B2 JP4177779A JP17777992A JP2622327B2 JP 2622327 B2 JP2622327 B2 JP 2622327B2 JP 4177779 A JP4177779 A JP 4177779A JP 17777992 A JP17777992 A JP 17777992A JP 2622327 B2 JP2622327 B2 JP 2622327B2
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nucleic acid
sequence
primer
dna
strand
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JPH05308971A (ja
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バンクス マリス カリー
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エフ．ホフマン−ラ ロシュ アクチェンゲゼルシャフト
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Description

【発明の詳細な説明】
【０００１】
【産業上の利用分野】本発明は、核酸配列を増幅するた
めの手段及びその使用に関する。より詳細には本発明
は、与えられたＤＮＡ又はＲＮＡ配列から初期に存在す
る量に比較してより大量の任意の特定の核酸配列を生成
せしめる方法に関する。該ＤＮＡ又はＲＮＡは単鎖又は
二重鎖であってもよく、比較的純粋であっても核酸の混
合物の一成分であってもよい。本発明の手段では、所望
の核酸配列の増幅を達成するために反応を繰り返し行う
ようにする。
【０００２】
【従来の技術】特に診断上の用途のためには、標的核酸
配列は問題のＤＮＡ又はＲＮＡのほんの僅かな部分であ
ることがあり、非同位体標識又は末端標識オリゴヌクレ
オチドプローブを使用するのではその存在を検出するこ
とは困難である。プローブ検出システムの感度を向上さ
せるために多くの労力が費やされているが、現在利用で
きる方法を用いて容易に検出できるに充分な量を得るた
めに、標的配列を増幅するような研究は殆ど行われてい
ない。核酸を初めから、あるいは既存の配列から合成す
る方法がいくつかの文献に記載されている。これらの方
法は、完全に特定された配列の与えられた核酸を大量に
生産することを可能にするものである。
【０００３】核酸を初めから合成する１つの既知方法
は、ヌクレオシド誘導体からの核酸の有機合成を含むも
のである。この合成は溶液中又は固体担体上で行われ
る。有機合成の１つのタイプはリン酸トリエステル法で
あり、これは遺伝子断片又は短い遺伝子を調製するため
に利用される。リン酸トリエステル法では、オリゴヌク
レオチドが調製され、次にこれは結合されてより長鎖の
核酸を形成する。この方法は、Ｓ．Ａ．ナーランクらに
より、Ｍｅｔｈ．Ｅｎｚｙｍｏｌ．６８巻９０頁（１９
７９年）及び米国特許第４，３５６，２７０号に開示さ
れている。該特許はソマトスタチン遺伝子の合成とクロ
ーニングを開示している。
【０００４】有機合成の第２のタイプはリン酸ジエステ
ル法であり、これはトランスファーＲＮＡ遺伝子の調製
に利用されている。この方法はＥ．Ｌ．ブラウンらによ
りＭｅｔｈ．Ｅｎｚｙｍｏｌ．６８巻１０９頁（１９７
９年）に開示されている。リン酸トリエステル法と同じ
ように、リン酸ジエステル法もオリゴヌクレオチドの合
成を含み、これらが実質的に結合されて所望の核酸が形
成される。
【０００５】上記した初めからの合成法は核酸の長鎖を
合成するために利用されるが、核酸を大量合成するため
の実用的方法ではない。両法とも労力と時間を消費し高
価な装置と試薬を必要としかつ全体収率が低い。全体収
率が低いのは、オリゴヌクレオチドの合成とそれらを結
合する反応が非効率的であることに起因する。長鎖の核
酸を合成する際あるいは短鎖の核酸を大量に合成する場
合でさえも、多くのオリゴヌクレオチドを合成し多くの
結合反応を行うことが要求される。従ってこれらの方法
は任意で所望の核酸を大量に合成するには実用的ではな
い。
【０００６】初めに存在する少量の核酸から大量の核酸
を生産する方法も存在する。これらの方法は好適な宿主
系内での核酸のクローニングを含み、ここでは所望の核
酸は宿主の形質変換に使用される好適なベクター中に挿
入される。宿主が培養されるとベクターが複製され、所
望の核酸のコピーが生産される。核酸断片のサブクロー
ニングについては、Ｔ．マニアチスらにより、コールド
・スプリング・ラボラトリーのＭｏｌｅｃｕｌａｒ Ｃ
ｌｏｎｉｎｇ ３９０−４０１頁（１９８２年）に簡単
に記述されている。この技術については米国特許第４，
４１６，９８８号及び４，４０３，０３６号にも記述さ
れている。
【０００７】米国特許第４，２９３，６５２号に記載さ
れている核酸の第３の合成法は、上述の有機合成と分子
クローニング法を合わせたものである。該法では、所望
の核酸配列を作り上げるのに必要な好適な数のオリゴヌ
クレオチドを初めに合成し、次にこれらを次の挿入の前
に増殖により増幅されるベクターに挿入する。
【０００８】
【発明が解決しようとする課題】本発明は、この分子ク
ローニング法に幾らかの類似性を有している。しかし本
発明はいかなる生物の繁殖も必要とせず、従って繁殖に
伴って起こり得る危険や不都合を回避することができ
る。本発明は所望の核酸と関連しない核酸の合成を必要
とせず、従って本発明によれば複雑な生物学的混合物か
らコストをかけて生成物を精製することも回避できる。
【０００９】本発明はプライマーと重合試薬を用いて１
種の核酸又は複数の核酸の混合物中に存在する１又は２
以上の特定の核酸配列を増幅する手段及びその手段の使
用に関し、増幅した配列は検出することができる。プラ
イマーは、そして１つのプライマーの伸長生成物は、他
のプライマーとハイブリダイズしたときに所望の特定の
核酸配列の生成のための鋳型となり、又その逆も起こ
る。そしてこのプロセスは所定量の配列が生成するまで
必要なだけ繰り返される。標的配列から大量の核酸を比
較的短時間で生産するためには、本方法は上記の従来法
より効率的であると期待される。本方法は核酸混合物に
僅かしか含まれていない核酸種を増幅し、該種を効率的
に検出するために特に有用である。
【００１０】
【課題を解決するための手段】本発明は特に、１種類の
核酸又は複数種類の混合物（各核酸は同一もしくは異る
長さの２本の別個の相補鎖から成るか又は１本鎖であ
る）中に含まれる少なくとも１つの核酸配列を指数的に
増幅するための手段であって、増幅されるべき各特定の
配列の異る鎖のそれぞれに実質的に相補的なプライマー
を提供するように、増幅されるべき各異る特定の核酸配
列のためのプライマーを含んで成る手段、及びその使用
に関する。
【００１１】１つのプライマーの伸長生成物は、他のプ
ライマーとハイブリダイズしたときに所望の特定の核酸
配列の生成のための鋳型となり、又その逆も起こる。そ
してこのプロセスは所定量の配列が生成するまで必要な
だけ繰り返される。標的配列から大量の核酸を比較的短
時間で生産するためには、本方法は上記の従来法より効
率的であると期待される。本方法は核酸混合物に僅かし
か含まれていない核酸種を増幅し、該種を効率的に検出
するために特に有用である。
【００１２】前記使用の１つの態様は、核酸又は核酸混
合物中に存在する少なくとも１種の特定の核酸配列を増
幅する方法であって、この場合、各核酸は同じ長さ又は
異る長さの２つの別個の相補的な鎖から成り、そして前
記の方法は、 （ａ）前記鎖を、各異る特定の配列を増幅するための２
つのオリゴヌクレオチドプライマーにより、増幅される
べき各異る配列について各核酸鎖に相補的な各プライマ
ーの伸長生成物が合成される条件下で処理し、各核酸鎖
に相補的な各プライマーの延長生成物が合成され、ここ
で、前記プライマーは各特定の配列の異る鎖と実質的に
相補的であって１方のプライマーから合成された延長生
成物がその相補体から分離された場合に他方のプライマ
ーの延長生成物の合成のためのプライマー鋳型として機
能することができるように選択され； （ｂ）前記プライマー延長生成物をそれらが合成された
鋳型から分離して単鎖分子を生成せしめ；そして （ｃ）段階（ｂ）から生じた単鎖分子を段階（ａ）のプ
ライマーにより、段階（ｂ）において生成した各単鎖を
鋳型としてプライマー延長生成物が合成される条件下で
処理する；ことを含んで成る。
【００１３】これらの段階は逐次的に又は同時に行うこ
とができる。さらに、段階（ｂ）及び（ｃ）は配列の所
望のレベルの増幅が得られるまで反復することができ
る。
【００１４】この発明は、完全に特定された配列の既存
の核酸を多量に製造するためのみならず、存在すること
は知られているがしかし完全には特定されていない核酸
配列を製造するためにも有用である。いずれの場合に
も、増幅されるべき配列の最初のコピーは入手可能でな
ければならない。但しそれは純粋である必要はなく、又
は別個の（ｄｉｓｃｒｅｔｅ）分子である必要はない。
【００１５】
【具体的な説明】プライマー、プローブ、検出すべきオ
リゴマー断片、オリゴマー対照体、及び標識されていな
いブロッキングオリゴマーに関して使用される「オリゴ
ヌクレオチド」という用語は、２又はそれ以上の好まし
くは３より多くのデオキシリボヌクレオチド又はリボヌ
クレオチドから成る分子として定義される。その正確な
大きさは多くの因子に依存し、その因子はオリゴヌクレ
オチドの究極的な機能と用途に依存する。
【００１６】ここで使用される「プライマー」という用
語は、精製された制限消化物として自然に存在しあるい
は合成的に調製されたオリゴヌクレオチドを意味し、こ
のプライマーは、例えば好適な温度及びpHでヌクレオチ
ドとＤＮＡポリメラーゼのような重合試薬が存在するよ
うな、核酸鎖に相補的なプライマーの伸長生成物の合成
が誘発される条件下に置かれたときに合成開始点として
機能することができる。
【００１７】該プライマーは増幅効率を最大にするため
単鎖であることが好ましいが、その代わりに二重鎖であ
ってもよい。二重鎖であると、プライマーは伸長生成物
を調製するために使用される前にまずその鎖を分離する
ために処理される。プライマーはオリゴデオキシリボオ
クレオチドであることが好ましい。プライマーは、重合
試薬の存在下で伸長生成物の合成を開始するために十分
な長さでなければならない。
【００１８】プライマーの正確な長さは、温度やプライ
マー源を含む多くの因子に依存する。例えば、目的とす
る配列の複雑さに依存してオリゴヌクレオチドプライマ
ーは典型的には１５から２５又はそれより多くのヌクレ
オチドを含むが、より少ないヌクレオチドを含むもので
あってもよい。短いプライマー分子は、鋳型とともに十
分安定なハイブリドの複合体を形成するために、より低
い温度を要求する。プライマーは増幅されるべき各特定
配列の異なる鎖と「実質的」に相補的であるように選択
される。このことはプライマーはそれぞれの鎖とハイブ
リダイズするに十分に相補的でなければならないことを
意味する。従ってプライマーの配列は鋳型の配列を正確
に反映する必要はない。
【００１９】例えば相補的でないヌクレオチド断片を、
プライマーの配列の残部が鎖に相補的であるようにプラ
イマーの５′末端に結合させてもよい。代わりに、プラ
イマーの配列が増幅されるべき鎖の配列と十分な相補性
を有していてそれらとハイブリダイズし、それによって
他方のプライマーの伸長生成物合成用鋳型を形成するな
らば、相補的でない塩基又はより長い配列がプライマー
内に散在してもよい。
【００２０】本発明で使用される「制限エンドヌクレア
ーゼ」及び「制限酵素」という用語は、二重鎖ＤＮＡを
特定の核酸配列又はその近傍で切断するような細菌性酵
素を意味する。本発明で使用される「ＤＮＡの多形現
象」という用語は、ＤＮＡ中の特定部位に２又はそれよ
り多くの異なったヌクレオチド配列が存在できる状態を
意味する。「制限断片長さの多形現象（ＲＡＬＰ）」と
いう用語は、特定の制限エンドヌクレアーゼによる消化
により形成される制限断片の長さに個体間の相違がある
ことを意味する。
【００２１】本発明は、核酸中に存在すると思われる１
又はそれ以上の所望の特定の核酸配列を増幅する方法に
関する。本法によれば大量の特定の配列を調製できるの
で、本発明はＤＮＡ又はメッセンジャーＲＮＡのクロー
ニング効率を改良し、かつ標的配列を増幅してその検出
を容易にするために使用することができる。この発明は
また、不完全な化学合成から生ずる核酸の混合物から所
望の配列を多量に得るためにも有用である。
【００２２】一般に、本発明の方法は、用いられる反応
ステップの数に関連して指数的な収量で少なくとも１つ
の特定の核酸配列を生産する連鎖反応を含み、該配列は
（ａ）必要とされる末端が、それとハイブリダイズする
オリゴヌクレオチドを合成できるに十分な程度に詳細に
知られており、（ｂ）連鎖反応を開始するために少量の
配列が入手可能であることが条件となる。連鎖反応で得
られる生成物は、使用した特定のプライマーの末端に対
応する末端を有するような別個の核酸のデュプレックス
である。
【００２３】精製された状態でも精製されていない状態
でもよい任意の核酸源を、所望の特定の核酸配列を含む
と思われるのであれば、出発核酸として使用できる。従
って本法では、例えば単鎖であっても二重鎖であっても
よいＤＮＡ又はＲＮＡ例えばメッセンジャーＲＮＡを使
用することができる。更にそれぞれ１つの鎖を含むＤＮ
Ａ−ＲＡＮハイブリドを使用してもよい。
【００２４】これらの核酸の混合物を使用してもよく、
又先行する増幅反応において同じか又は異なったプライ
マーを用いて生産された核酸を使用してもよい。増幅す
べき特定の核酸配列は大きな分子の一部であってもよ
く、特定の配列が核酸全体を構成するようにはじめから
個別的な分子として存在していてもよい。増幅すべき配
列は初めから純粋な状態で存在する必要はなく、該配列
は、複雑な混合物の小部分、例えば全ヒトＤＮＡ中のβ
−グロビン遺伝子、又は特定の生物的試料の極く僅かの
部分のみを構成する特定の微生物に起因する核酸配列の
部分であってもよい。
【００２５】出発物質としての核酸は、同じか又は異な
った２以上の所望の特定の核酸配列を含んでいてもよ
い。従って本発明の方法は、１つの特定の核酸配列を大
量に生産するだけでなく、同じか又は異なった核酸分子
上に位置する２以上の異なった特定の核酸配列の同時増
幅にも有用である。
【００２６】核酸は、例えばｐＢＲ３２２のようなプラ
スミド、クローン化されたＤＮＡ又はＲＮＡ、又は細
菌、酵母、ビールス及び植物や動物などの高級生物等の
自然にあるＤＮＡ又はＲＮＡ等の任意源から得ることが
できる。ＤＮＡ又はＲＮＡは、例えばマニアチスらによ
りＭｏｌｅｃｕｌａｒ Ｃｌｏｎｉｎｇの２８０から２
８１頁（１９８２年）に記載されているような種々の技
術により、血や絨毛又は羊膜細胞等の組織物質から抽出
することができる。
【００２７】本発明方法により、任意の特定の核酸配列
を生産することができる。配列の両末端の十分な数の塩
基が十分詳細に分かっており、これにより所望の配列の
異なる複数の鎖に対し、かつ該配列に沿った次のような
相対位置にハイブリダイズする２つのオリゴヌクレオチ
ドプライマーを調製することができればよく、すなわ
ち、１つのプライマーから合成伸長した生成物が、鋳型
（相補体）から分離されたときに、限定された長さの核
酸中へ他のプライマーを伸長させるための鋳型としての
役割を果たせばよい。
【００２８】配列の両末端の塩基に関する知識が増加す
るほど目的とする核酸配列のためのプライマーの特異性
も大きくなり、従って本法の有効性も大きくなる。以後
使用するプライマーという用語は、特に増幅すべき断片
の末端配列に関する情報にいくらかの曖昧さがある場合
には、１より大きい数のプライマーを意味するものと理
解されるべきである。例えば、核酸配列が蛋白質配列の
情報から推測できる場合、遺伝子コードの縮重に起因す
るすべての可能なコドン変化を示す配列を含むプライマ
ーを集めて各鎖用として使用する。このような集合のう
ちの１つのプライマーは、増幅すべき所望配列の末端と
一致する。
【００２９】オリゴヌクレオチドプライマーは任意の好
適な方法、例えば上記したリン酸トリエステル法及びリ
ン酸ジエステル法又はそれらのオートメーション化され
た方法を使用して調製することができる。このようなオ
ートメーション化された方法のうちの１つによれば、ビ
ューケージらによりＴｅｔｒａｈｅｄｒｏｎ Ｌｅｔｔ
ｅｒｓ２２巻１８５９−１８６２頁に記載されている通
り、ジエチルフォスフォロアミダイトを出発物質として
使用して合成することができる。修飾された固体担体上
でのオリゴヌクレオチド合成の１つの方法が米国特許第
４，４５８，０６６号に記載されている。生物源（例え
ば制限エンドヌクレアーゼ消化物）から分離したプライ
マーを使用することも可能である。
【００３０】特定の核酸配列は、該配列を鋳型として含
む核酸を使用して生産される。核酸が２つの鎖を含んで
いるときは、別のステップとしてでもプライマーの伸長
生成物の合成と同時じもよいが、該核酸は鋳型として使
用される前に鎖を分離する必要がある。この鎖分離は、
物理的、化学的及び酵素的方法を含む任意の好適な変性
法により行うことができる。核酸の鎖を分離する１つの
物理的方法は、完全に（９９％以上）変性されるまで核
酸を加熱することを含む。
【００３１】典型的な加熱変性は８０から１５０℃で１
から１０分間加熱することを含む。鎖の分離は、ヘリカ
ーゼ、又はヘリカーゼ活性を有し、リボＡＴＰの存在下
でＤＮＡを変性させるものとして知られる酵素ＲｅｃＡ
として知られる酵素類からの１酵素により誘発させるこ
ともできる。ヘリカーゼで核酸の鎖を分離するのに好適
な反応条件はクーン ホフマン ベーリングによりＣＳ
Ｈ−Ｑｕａｎｔｉｔａｔｉｖｅ Ｂｉｏｌｏｇｙの４３
から６３頁（１９７８年）に記載され、ＲｅｃＡを使用
する技術は、Ｃ．ラディングによりＡｎｎ．Ｒｅｖ．Ｇ
ｅｎｅｔｉｃｓの１６巻４０５から４３７頁に記載され
ている。
【００３２】増幅すべき配列を含む当初の核酸が単鎖で
あると、その相補体をそれに１つ又は２つのオリゴヌク
レオチドプライマーを加えて合成する。好適な単一プラ
イマーが加えられると、プライマー、重合試薬及び後述
する４つのヌクレオチドの存在下でプライマーの伸長生
成物が合成される。生成物は部分的に単鎖の核酸と相補
的で、核酸鎖とハイブリダイズして長さの異なるデュプ
レックスを形成し、これは上記した通り単鎖に分離さ
れ、相補的な２つの分離された鎖となる。代わりに２つ
の好適なプライマーを単鎖に加えて反応を行うこともで
きる。
【００３３】当初の核酸が増幅すべき配列を構成するな
らば、プライマーの伸長生成物は当初の核酸の鎖と完全
に相補的となり、ハイブリダイズして同じ長さの鎖から
成るデュプレックスを形成し、これは分離されて単鎖の
分子となる。核酸の相補的な鎖が分離すると、当初の核
酸が二重鎖であっても単鎖であっても、その鎖は他の核
酸鎖の合成用鋳型として容易に使用することができる。
この合成は任意の好適な方法を用いて行うことができ
る。通常それは好ましくはpHが７から９、最も好ましく
は８である緩衝水溶液中で起こる。好ましくは過剰のモ
ル比（クローン化された核酸については、通常プライマ
ー対鋳型が１０００：１、そしてゲノムの核酸について
は通常プライマー対鋳型が１０6 ：１）の２つのオリゴ
ヌクレオチドプライマーを分離された鋳型鎖を含む緩衝
水溶液中に加える。
【００３４】しかし本法を診断的用途に使用する場合に
は相補的な鎖の量は既知ではないことを理解すべきであ
り、従って相補的な鎖の量に関連するプライマーの量を
確信をもって決定することはできない。しかし実際に
は、増幅すべき配列が複雑な長鎖の核酸鎖の混合物中に
含まれる場合には、加えられるプライマーの量は相補的
な鎖（鋳型）の量よりも通常モル過剰とする。本法の効
率を改良するためには、大きな過剰モル比とすることが
好ましい。
【００３５】デオキシリボヌクレオシド三リン酸である
デオキシＡＴＰ、デオキシＣＴＰ、デオキシＧＴＰ及び
ＴＴＰの十分な量も合成混合物に加え、生成する溶液を
約９０−１００℃で約１から１０分、好ましくは１から
４分間加熱する。この加熱時間の後、溶液の温度をプラ
イマーのハイブリダイゼーションに好適な室温まで下げ
る。
【００３６】この冷却した混合物にプライマー伸長反応
を誘導し又は触媒するための適当な薬剤（誘導試薬又は
重合試薬と称する）を加え、従来知られている条件下で
反応を行わせる。この合成反応は、室温からそれを越え
ると重合試薬が効率的に機能しない温度までの間で行わ
せることができる。従って例えばＤＮＡポリメラーゼを
重合試薬として使用するときは、温度を通常４０℃以上
に上昇させない。最も好ましくは反応は常温において起
こる。
【００３７】重合試薬（誘導試薬）は、プライマーの伸
長生成物の合成を達成できるものならば、酵素を含むど
のような化合物でも系でもよい。この目的のための好適
な酵素は、例えばＥ．コーリーＤＮＡポリメラーゼＩ、
Ｅ．コーリーＤＮＡポリメラーゼＩのクレノー断片、Ｔ
４ＤＮＡポリメラーゼ、他の入手できるＤＮＡポリメラ
ーゼ、逆転写酵素及び耐熱性酵素を含む他の酵素を含
み、これらは好適な態様でヌクレオチドの結合を促進
し、各核酸鎖と相補的であるプライマーの伸長生成物を
形成する。
【００３８】一般に合成は各プライマーの３′末端から
始まり、合成が終了するまで鋳型鎖に沿って５′末端方
向に向かって進行し、異なった長さの分子を生成する。
しかし上述の方法と同じ方法を用いて５′末端で合成を
始め、他の方向に向かって反応を進行させる試薬があろ
う。新たに合成された鎖とそれと相補性を有する核酸鎖
は、本法のその後のステップにおいて使用される二重鎖
分子を形成する。次のステップでは、二重鎖分子の鎖は
上述の任意の手順を用いて分離され、単鎖分子を提供す
る。
【００３９】新たな核酸が該単鎖分子上で合成される。
追加の誘導試薬、ヌクレオチド及びプライマーを、上記
に規定した条件下で反応を進行させるために必要ならば
加えてもよい。オリゴヌクレオチドプライマーの一末端
から再度合成が始まり、そして鋳型の単鎖に沿って進行
して他の核酸を生成する。このステップの後における伸
長生成物の半分は２つのプライマーが結合した特定の核
酸配列から成っている。鎖分離と伸長生成物合成のステ
ップは、特定の核酸配列を所定量生産するまで必要なだ
けの回数繰り返すことができる。後により詳細に記載す
るように、特定の核酸配列は指数的に蓄積する。
【００４０】最初の核酸又は核酸の混合物から２以上の
特定の核酸配列を生産することが望ましい場合は、好適
な数の異なったオリゴヌクレオチドプライマーを使用す
る。例えば２つの異なった特定の核酸配列を生成する場
合には、４つのプライマーを使用する。プライマーのう
ちの２つは特定の核酸配列のうちの１つに関するもの
で、他の２つのプライマーは第２の特定の核酸配列に関
するものである。これにより、２つの異なった特定の配
列が本法を用いて指数的に生産され得る。本発明は、各
ステップ後に新しい試薬を加える段階的方法、又は全て
の試薬を初期のステップで加える同時的方法、又はある
与えられた数のステップの後に新しい試薬を加える一部
段階的で一部同時的である方法のいずれによっても行う
ことができる。
【００４１】熱処理のように重合試薬を不活性化する鎖
分離方法を採用した場合には、熱に対して不安定である
酵素の場合がそうであるように、各鎖分離ステップ後に
重合試薬を補充することが必要である。ヘリカーゼのよ
うな酵素的手段を含む多数の精製された成分を鎖分離ス
テップで使用する場合は、同時的方法を使用することが
できる。
【００４２】同時的方法では、反応混合物は、所望の配
列を含む核酸鎖の他に、鎖分離酵素（例えばヘリカー
ゼ）、ｒＡＴＰのような鎖分離酵素への適切なエネルギ
ー供給源、４つのヌクレオチド、モル過剰のオリゴヌク
レオチドプライマー及びＥ．コーリーＤＮＡポリメラー
ゼＩのクレノー断片のような誘導試薬を含むことができ
る。同時的方法で変性のために熱を使用するときは、誘
導試薬に依存するが好ましくは ６５−９０℃の高温で
機能する熱安定性ポリメラーゼ等の熱安定性誘導試薬を
使用し、この温度で核酸は平衡状態にある単鎖と二重鎖
から成っている。
【００４３】長さの短い核酸には、約５０℃程度の低温
が採用される。どの程度の高温が使用できるかは、その
温度で酵素が失活するかあるいはプライマーのハイブリ
ダイゼーションが不十分な程度しか起こらないかどうか
に依存する。このような熱安定性酵素は、例えばＡ．
Ｓ．カレディンらによりＢｉｏｋｈｉｍｉｙａ４５巻６
４４−６５１頁（１９８０年）に記載されている。本法
の各ステップは、全ての試薬が始めから存在するにもか
かわらず、続いて起こる。必要ならば追加の試薬を加え
てもよい。適切な長さの時間が経過して所望量の特定の
核酸配列が生成した後、任意の公知方法で酵素を失活さ
せるか反応成分を分離するかして反応を停止させる。
【００４４】本発明方法は連続的に行ってもよい。オー
トメーション化された方法の一態様として、反応を、変
性区域、試薬添加区域及び反応区域を通ってサイクルさ
せるような方法がある。他の態様では、プライマーの伸
長生成物の合成に使用する酵素をカラム中で固定化する
ことができる。他の反応成分は連続するカラムと加熱用
コイルを通るようにポンプを使って連続的に循環され、
これにより、生成した核酸が酵素を失活させることなく
繰り返し変性される。
【００４５】本発明の概略が下記に示され、ここでは相
補的な鎖〔Ｓ+ 〕と〔Ｓ- 〕から成る所望配列〔Ｓ〕を
含む二重鎖ＤＮＡが核酸として使用されている。第１の
及び引き続いて起こる反応サイクルでは、当初の鋳型上
の各オリゴヌクレオチドプライマーの伸長は、プライマ
ーの１つとともにのみ終了する制限のない長さの、１つ
の新しいｓｓＤＮＡ分子生成物を生成する。以後「長鎖
生成物」と呼ぶこれらの生成物は直線的に蓄積し、つま
り任意数のサイクルの後に存在する量はサイクル数に比
例する。
【００４６】このように生産される長鎖生成物は、引き
続いて起こるサイクルの間一方又は他方のオリゴヌクレ
オチドプライマーの鋳型として機能し、所望配列
〔Ｓ+ 〕又は〔Ｓ- 〕の分子を生成する。これらの分子
も一方又は他方のオリゴヌクレオチドプライマーの鋳型
として機能し、更に他の〔Ｓ+ 〕及び〔Ｓ- 〕を生成
し、従ってサイクル数に関連して指数的に〔Ｓ〕の蓄積
を生じさせる連鎖反応が維持される。オリゴヌクレオチ
ドのハイブリダイゼーションにより形成される意図され
ない副生成物は、それ自身触媒活性がなく（稀な例を除
く）、従って直線的に蓄積する。
【００４７】
【表１】
【００４８】
【表２】
【００４９】
【表３】
【００５０】
【表４】
【００５１】一方のプライマーのオリゴヌクレオチド配
列と共に終る鎖及び他方の相補的配列の各鎖は、生産す
ることが望まれている特定の核酸配列〔Ｓ〕であること
が分かる。本法のステップは、プライマー１及び２、重
合試薬及び存在するヌクレオチドの量によってのみ限定
される以外、無限に繰り返すことができる。最初の核酸
は複製されないので、その量は全工程中一定である。長
鎖生成物は最初の核酸からのみ生産されるので、その量
は直線的に増加する。特定の配列の量は指数的に増加す
る。従って特定の配列はその量が増加して優勢な成分と
なる。このことは次表に示され、該表は各サイクルの効
率が１００％であるとした場合のｎサイクル後の理論的
に存在する成分量を比較したものである。
【００５２】 ０からｎサイクル後の二重鎖の数 サイクル数 鋳型 長鎖生成物 特定配列〔Ｓ〕 ０ １ −− −− １ １ １ ０ ２ １ ２ １ ３ １ ３ ４ ５ １ ５ ２６ １０ １ １０ １，０１３ １５ １ １５ ３２，７５２ ２０ １ ２０ １，０４８，５５５ ｎ １ ｎ ２n −ｎ−１ 単鎖の核酸を鋳型として使用すると、サイクルあたり１
つの長鎖生成物が生成する。
【００５３】更に本法はインビトロの突然変異用として
使用することができる。オリゴデオキシリボヌクレオチ
ドは増幅されるべきＤＮＡ配列と正確に相補的である必
要はない。これらは、ポリメラーゼ酵素や他に使用され
るいずれかの誘導試薬によって伸長されるために十分な
程度に、鎖とハイブリダイズすることができればよい。
使用するプライマーが最初の鋳型と正確に相補的でない
場合のポリメラーゼ連鎖反応の生成物は鋳型よりむしろ
プライマー配列を有し、これによりインビトロの突然変
異を可能にする。
【００５４】引き続くサイクルでは、より以上のミスペ
ア−プライミングが必要とされないので、この突然変異
が減ることのない効率下で増幅される。このように生産
された突然変異体は標準的な分子生物学的技術により適
切なベクター中へ挿入され、変化した蛋白質を生産する
能力等の変化した特質をこのベクターに与える。
【００５５】上述した変化したＤＮＡ配列を形成する方
法は、より以上の配列変化を誘発させるために異なった
プライマーを使用して該変化したＤＮＡに対して繰り返
すことができる。この方法では、一連の突然変異配列が
徐々に生成され、ここで、この一連のものに新しく加え
られるものは、最後のものと僅かに異なることができる
が、最初のＤＮＡ源配列とは非常に大きく異なることが
できる。この方法では、非常に大きなミスマッチの場合
にプライマーが機能しないために単一ステップでは行う
ことのできない変化を、最終的には作り出すことができ
る。
【００５６】更に、十分な量のプライマーが増幅される
鎖に相補的である配列を含むのであれば、プライマーは
その配列の一部として相補的でない配列を含むことがで
きる。例えば鋳型配列に相補的でない核酸配列（例えば
プロモーター、リンカー、コード配列等）を、１つ又は
両方のプライマーの５′末端に結合させることができ、
これにより増幅工程の生成物にこれを付加することがで
きる。
【００５７】伸長プライマーを添加した後、相補的でな
い核酸挿入部を含む新しい鋳型の所望量を得るために十
分な数のサイクルを実施する。これにより簡単な技術を
用いて比較的短時間（例えば２時間又はそれ以下）内に
組合わされた断片を大量に生産することが可能になる。
【００５８】本法は、伝染性疾患、遺伝性疾患又は細胞
性疾患、例えば癌、と関連する特定の核酸配列、例えば
発癌遺伝子、の検出及び／又は特徴付けを可能にするた
めに使用される。増幅は、例えば胎児細胞から得られる
ＤＮＡを用いる鎌状赤血球貧血の胎児診断等、分析に利
用できる核酸の量が非常に小さい場合に有用である。増
幅は、本来的に感度の良くない非放射性検出技術を用い
て少量の試料を分析する場合、又は放射性技術を用いる
が迅速な検出が望ましい場合に特に有用である。
【００５９】本発明の目的のためには、遺伝子疾患は、
例えば鎌状赤血球貧血、嚢胞性繊維症、α−サラセミ
ア、β−サラセミア等の、任意の生物体からのゲノムＤ
ＮＡ中の特定の欠損及び／又は変異を含む。鎌状赤血球
貧血は本法による好適なＤＮＡ配列の増幅の後のオリゴ
マー制限分析又はＲＦＬＰ状分析を経て容易に検出する
ことができる。α−サラセミアは配列が存在しないこと
により検出することができ、β−サラセミアは疾患を起
こさせる変異に近接してリンクする多形性（ｐｏｌｙｍ
ｏｒｐｌｉｃ）制限部位の存在により検出することがで
きる。
【００６０】これら全ての遺伝子疾患は適切な配列を増
幅し、それを放射性プローブを使用せずにサザンプロッ
ト法により分析して検出することができる。このような
方法では、例えば非常に少量の所望配列を含む羊水から
のＤＮＡの少量の試料を増幅し、制限酵素で切断し、そ
してサザンブロット法で分析する。増幅シグナルをハイ
レベルとすることにより、非放射性標体を使用すること
が容易になる。
【００６１】他の態様では、少量のＤＮＡを便利なレベ
ルまで増幅し、次に更に伸長反応を行うが、この場合容
易に検出できるヌクレオチド誘導体（例えば32Ｐ又はビ
オチンでラベルしたヌクレオチド三リン酸）を直接最終
のＤＮＡ生成物に導入し、これを制限分析及び電気泳動
分析あるいは任意の他の好適な方法を用いて分析する。
この技術のモデル系の例を図１２に示してある。
【００６２】図３のモデル系に示した更に他の態様で
は、核酸は増幅の前に特定の制限エンドヌクレアーゼに
暴露する。切断された配列は増幅できないので、予め制
限酵素で処理したＤＮＡ試料の存在にもかかわらず増幅
された断片が現れることは増幅された配列中にエンドヌ
クレアーゼの部位がないことを暗示する。増幅された配
列が存在するか否かは適当な方法で検出することができ
る。
【００６３】この技術の実際的な適用方法は、本明細書
とサイキらによるＢｉｏｔｅｃｈｎｏｌｏｇｙ３巻１０
０８−１０１２頁に記載されているオリゴマー制限技術
を用いて鎌状赤血球貧血の検出を容易にする使用により
例示することができる。鎌状赤血球貧血はβ−グロビン
遺伝子の第６コドンの１つの塩基対の変化により生ずる
ヘモグロビンの疾患である。
【００６４】図１３は多形現象（ｐｏｌｙｍｏｒｐｈｉ
ｓｎ）領域中の正常及び鎌状赤血球貧血のβ−グロビン
遺伝子の配列を示すもので、一本線は正常遺伝子にのみ
存在するＤｄｅＩ部位の位置を示し、二本線は正常及び
鎌状赤血球貧血対立遺伝子の両方に存在する非多形性の
ＨｉｎｆＩ部位の位置を示す。図１４は両制限部位部位
間にわたり星印で示された部分がラベルされているプロ
ーブを用いて正常のβ−グロビンＤＮＡをオリゴマー制
限開裂する方法を示すものである。
【００６５】前に記載したようにして増幅されたＤＮＡ
が変性され、ラベルされたプローブとアニールされる。
酵素ＤｄｅＩはＤＮＡを再構成されたＤｄｅＩ部位で開
裂させ、ラベルされたオクタマーを生じさせる。テスト
に使用した条件下では、オクタマーはデュプレックスか
ら離れるのに十分な短さである。引き続く酵素Ｈｉｎｆ
Ｉの添加は今や単鎖であるオクタマーに何の影響も与え
ない。図１５はβ−グロビンＤＮＡの鎌状赤血球対立遺
伝子に適用した前記と同じ方法を示す。酵素ＤｄｅＩ
は、Ａ−Ａの塩基対がミスマッチしたものであるため、
増幅されたＤＮＡとラベルされたプローブとで形成され
たデュプレックスを開裂させることはできない。
【００６６】しかし酵素ＨｉｎｆＩはハイブリッド制限
開裂せしめ、ラベルされたトリマーが生成される。実際
にはこの方法は、特定のシグナルがいずれかの対立遺伝
子の存在と関連するので、個体のＤＮＡが野性型のホモ
接合体か、鎌状赤血球貧血型のホモ接合体か又は鎌状赤
血球貧血形質を有するヘテロ接合体であるかを検診する
ことができる。上述の方法を使用して適切な配列を増幅
させることにより１つの32Ｐラベルのみを有するプロー
ブを用いて単コピー遺伝子を迅速に分析することができ
る。
【００６７】種々の伝染性疾患は、原因となる微生物に
特異的である特定のＤＮＡ配列の臨床試料中での存在に
より診断することができる。これらはサルモネラ、クラ
ミジア、ネイセリア等の細菌、肝炎ビールス等のビール
ス、マラリアの原因となるプラスモジウム（Ｐｌａｓｍ
ｏｄｉｕｍ）等の寄生体を含む。ファルコーに与えられ
た米国特許第４，３５８，５３５号は、伝染性疾患の診
断用の特別なＤＮＡハイブリダイゼーションプローブの
使用につき記述している。
【００６８】ファルコー法に固有の問題は、感染した患
者からの臨床試料中には比較的少ない数の病原生物しか
存在せず、これらから抽出されたＤＮＡは試料中の全Ｄ
ＮＡの非常に小さな部分を構成するのみであるというこ
とである。ＤＮＡ試料を固定化しハイブリダイゼーショ
ン検出する前に問題となっている配列を特異的に増幅す
ることは、これらの方法の感度と特異性を大きく改良す
る。
【００６９】伝染性疾患の診断用にＤＮＡプローブを臨
床的にルーチン化して使用することは、ワードのヨーロ
ッパ特許第６３，８７９号に記載されたいるように非放
射的にラベルされたプローブを使用するのならば、大い
に簡略化される。この方法では、ビオチンを含むＤＮＡ
プローブがアビジン又はビオチンに特異的な抗体に結合
した色素体（ｃｈｒｏｍｏｇｅｎｉｃ）酵素により検出
される。この型の検出は便利であるが、比較的低感度で
ある。本法による特異的なＤＮＡ増幅と安定にラベルさ
れたプローブを組み合わせることにより、ファルコー及
びワードの方法をルーチン化した臨床における有用な方
法に実施するのに要求される便利さと感度を提供するこ
とができる。
【００７０】この増幅工程は単一コピーのヒト遺伝子か
ら十分な量のＤＮＡを調製するのに利用することもで
き、これにより臭化エチジウムのような簡単な非特異的
なＤＮＡ染色によりそれを検出でき、直接ＤＮＡ診断を
行うことができる。伝染性疾患及び生物体のゲノム中の
病原的異常性を検出するほか、本法は任意の病原状態と
関連しないＤＮＡ多形現象（ポルモルフィズム）を検出
するために使用することもできる。
【００７１】次の実施例は例示のために提示するもの
で、どのようにも本発明を限定することを意図するもの
ではない。これらの実施例で全てのパーセントは固体の
場合は重量で、液体の場合は容量であり、他に指定がな
い限り温度は摂氏温度である。
【００７２】実施例１ 次のヌクレオチド配列を有する２５塩基対配列 ５′CCTCGGCACCGTCACCCTGGATGCT ３′ ３′GGAGCCGTGGCAGTGGGACCTACGA ５′ （ＡＴＣＣから得られるｐＢＲ３２２の４７塩基対Ｆｏ
ｋＩ制限断片上に含まれる）を次のように調製した。４
７塩基対断片を含むｐＢＲ３２２のＦｏｋＩ消化物を供
給者であるニューイングランド社の指示による条件に従
ってｐＢＲ３２２をＦｏｋＩで消化することにより調製
した。使用したプライマーは、５′d(CCTCGGCACCG)３′
と５′d(AGCATCCAGGGTG)３′であり、通常の技術により
調製した。２５ｍＭのリン酸カリウムと１０ｍＭの塩化
マグネシウム、及び１００ｍＭの塩化ナトリウムから成
る緩衝液（pH７．５）３３μｌに２４３３ピコモルの上
述の各プライマー、２．４ピコモルのｐＢＲ３２２のＦ
ｏｋＩ消化物、２２ナノモルのデオキシＡＴＰ、２２ナ
ノモルのデオキシＣＴＰ：１９ナノモルのデオキシＧＴ
Ｐ及び１０ナノモルのＴＴＰを加えた。
【００７３】混合物を８５℃で５分間加熱し、室温まで
冷却した。Ｅ．コーリーＤＮＡポリメラーゼＩのクレノ
ー断片の５単位を加え、温度を１５分間維持した。その
後再度８５℃で５分間加熱し、冷却した。クレノー断片
の５単位を再度加え、１５分間反応を行った。加熱、冷
却及び反応の各ステップを更に１１回繰り返した。
【００７４】最後の繰り返しの後、５μｌを反応混合物
から取り出した。これを８５℃で３分間加熱し、室温に
冷却した。１２．５ピコモルのα−Ｐ32−デオキシシチ
ジン三リン酸と５単位のクレノー断片を加え反応を１５
分間進行させた。ラベルされた生成物をポリアクリルア
ミドゲルの電気泳動で確認した。１３サイクル後に見え
る強くラベルされたバンドのみが、意図する２５塩基対
配列であった。
【００７５】実施例２ 増幅されるべき所望の配列は、ヒトのβ−グロビン遺伝
子に含まれかつ鎌状赤血球貧血に関するＭｓｔII部位に
伸びる９４塩基対の配列であった。該配列は図１に示す
ヌクレオチド配列を有している。
【００７６】Ｉ．プライマーの合成 次の２つのオリゴデオキシリボヌクレオチドプライマー
を下記する方法を用いて調製した。 ５′CACAGGGCAGTAACG ３′プライマーＡ、及び ３′TTTGCTTCTGACACA ３′プライマーＢ
【００７７】オートメーション化された合成法 ビューケージとカルサース法（Ｔｅｔｒａｈｅｄｒｏｎ
Ｌｅｔｔｅｒｓ２２巻１８５９−１８６２頁（１９８
１年）に従って合成したジエチルフォスフォロアミダイ
トを、バイオサーチＳＡＭ−１を使用して制御した多孔
ガラス担体から誘導したヌクレオシドへ次々と濃縮し
た。この方法は、ジクロルメタン中でのトリクロル酢酸
による脱トリチル化と活性のある水素供与体としてベン
ゾトリアゾールを使用する縮合、及びテトラハイドロフ
ラン及びピリジン中での無水酢酸とジメチルアミノピリ
ジンによるキャッピングを含んでいた。１サイクルの時
間は約３０分であった。各ステップの収率は実質的に当
量的であり、脱トリチル化の間に解離するジメトキシト
リチルアルコールを集め分光器による検査で決定した。
【００７８】オリゴデオキシリボヌクレオシドを脱保護
化し、精製する方法 固体担体をカラムから取り出し、１mlの濃水酸化アンモ
ニウムに閉鎖管中室温で４時間曝した。担体を濾過で取
り除き、一部が保護されたオリゴデオキシリボヌクレオ
チドを含む溶液の温度を５５℃に上昇させ、５時間維持
した。アンモニアを取り除き、残渣を調製用ポリアクリ
ルアミドゲルに適用した。３０ボルト／cmで９０分間電
気泳動を行い、生成物を含むバンドを螢光プレート上の
ＵＶシャドウイングで同定した。
【００７９】該バンドを切り取り、１mlの蒸留水で一晩
かけて４℃で溶出した。この溶液をアルテックＲＰ１８
カラムにかけ、pH６．０の１％酢酸アンモニウム緩衝液
中７−１３％のアセトニトリルで溶出した。この溶出液
は２６０nmの紫外吸収でモニターし、適切なフラクショ
ンを集め、固定した量での紫外吸収で定量分析し、かつ
室温下で減圧遠心機中で蒸発させ乾燥した。
【００８０】オリゴデオキシリボヌクレオチドの特徴付
け 精製したオリゴヌクレオチドのテスト溶液をポリヌクレ
オチドキナーゼ及びγ32Ｐ−ＡＴＰで32Ｐラベルした。
このラベルした化合物を５０ボルト／cmで４５分間電気
泳動にかけた後、１４−２０％のポリアクリルアミドゲ
ルのオートラジオグラフィーで確認した。この方法では
分子量を確認することができる。ヘビ毒ジエステラーゼ
と細菌性アルカリフォスファターゼを使用してオリゴデ
オキシリボヌクレオチドをヌクレオシドに消化し、そし
て次に、逆相ＨＰＬＣカラム、並びに１０％アクリロニ
トリル及び１％酢酸アンモニウム移動相を使用して、誘
導されたヌクレオシドを分離し定量することにより塩基
組成を決定した。
【００８１】II．ＤＮＡ源 Ａ．全ヒト野性型ＤＮＡの抽出 正常のβ−グロビンのヒトゲノムＤＮＡホモ接合体を、
ステットラーらによりＰｒｏｃ．Ｎａｔ．Ａｃａｄ．Ｓ
ｃｉ．の７２巻５９６６−５９７０頁に記載された技術
を用いてセルラインＭｏｌｔ４（ヒューマン・ジェネテ
ィック・ミュータント・セル・レポジトリーから入手
し、ＧＭ２２１９ｃと同定した）から抽出した。
【００８２】Ｂ．クローン化したグロビン遺伝子の造成 正常のβ−グロビン遺伝子の１．９kbのＢａｍＨＩ断片
をコスミドｐＦＣ１１から分離し、ｐＢＡ３２２のＢａ
ｍＨＩ部位に挿入した（ソベロンらのＧｅｎｅ９巻２８
７−３０５頁（１９８０年））。合成４０塩基対プロー
ブとハイブリダイズする領域を含むこの断片は、第１及
び第２のエクソン、第２のイントロン、並びに遺伝子の
５′のフランキング（ｆｌａｎｋｉｎｇ）配列を含む
（ローンらのＣｅｌｌ１５巻１１５７−１１７４頁）。
このクローンはｐＢＲ３２８：ＨｂＡと名付けられ、Ａ
ＴＣＣ第３９，６９８号として１９８４年５月２５日に
寄託された。
【００８３】β−グロビンの鎌状赤血球貧血対立遺伝子
の対応する１．９kbのＢａｍＨＩ断片はコスミドｐＦ１
２から分離され、上述の通りクローン化された。このク
ローンはｐＢＲ３２８：Ｈｂｓと名付けられ、ＡＴＣＣ
第３９，６９９号として１９８４年５月２５日に寄託さ
れた。各組み換えプラスミドをＥ．コーリーＭＭ２９４
（ＡＴＣＣ第３９，６０７号）へ形質変換し、そして増
殖せしめた。
【００８４】Ｃ．クローン化されたグロビン遺伝子のＭ
ｓｔIIによる消化 それぞれの全量が１００μｇであるｐＢＲ３２８：Ｈｂ
ＡとＰｂｒ３２８：ＨｂＳを単独で２０単位のＭｓｔII
（ニューイングランドバイオラブ社）とともに１６時間
３７℃、１５０ｍＭＮａＣｌ，１２ｍＭのＴｒｉｓ Ｈ
Ｃｌ（pH７．５），１２ｍＭのＭｇＣｌ2 ，１ｍＭのジ
チオスレイトール（ＤＴＴ）及び１００μｇ／mlのウシ
血清アルブミン（ＢＳＡ）中で消化した。生成物はそれ
ぞれｐＢＲ３２８：ＨｂＡ／ＭｓｔII及びとｐＢＲ３２
８：ＨｂＳ／ＭｓｔIIと名付ける。
【００８５】III ．ポリメラーゼの連鎖反応 ６０ｍＭ酢酸ナトリウム、３０ｍＭトリス−アセテート
及び１０ｍＭ酢酸マグネシウムを含むpH８．０の緩衝液
１００μｌへ１００ピコモルのプライマーＡ（d(CACAGG
GCACTAACG)の配列）、１００ピコモルのプライマーＢ
（d(TTTGCTTCTGACACA)の配列）及び１０００ピコモルの
デオキシＡＴＰ、デオキシＣＴＰ、デオキシＧＴＰ及び
ＴＴＰを含む２μｌの溶液を加えた。更に上述した、下
記のＤＮＡ源の１つを加えた。
【００８６】１０μｇの全ヒト野性型ＤＮＡ（反応Ｉ） ０．１ピコモルのｐＢＲ３２８：ＨｂＡ（反応II） ０．１ピコモルのｐＢＲ３２８：ＨｂＳ（反応III ） ０．１ピコモルのｐＢＲ３２８：ＨｂＡ／ＭｓｔII（反
応IV） ０．１ピコモルのｐＢＲ３２８：ＨｂＢ／ＭｓｔII（反
応Ｖ） 非標的ＤＮＡ（反応VI）
【００８７】得られる各溶液を１００℃で４分間加熱し
２分間で室温まで冷却し、その後Ｅ．コーリーＤＮＡポ
リメラーゼのクレノー断片の４単位を含む１μｌを加え
た。各反応は１０分間行い、その後プライマー、ヌクレ
オチド及びＤＮＡを加え、加熱し、冷却し、ポリメラー
ゼを加え、そして反応させるサイクルを反応Ｉについて
は１９回、反応II−VIについては４回繰り返した。
【００８８】第１サイクルの前及び各反応の最後のサイ
クルの後で取り出された反応Ｉ及びIIのアリコート４マ
イクロリットルをpH８．３の０．０８９Ｍトリス硼酸塩
緩衝液中で、２．５ｍＭＥＤＴＡ中で、１２％ポリアク
リルアミドゲルに加えた。このゲルを２５ボルト／cm、
４時間電気泳動させ、固相担体として機能するナイロン
膜へ移し、そしてpH７．４で３０％のフォルムアミド、
３ｘＳＳＰＥ，５ｘデンハルツ及び５％のドデシル硫酸
ナトリウム中で、標準的技術を用いて調製した次式： ５′d(TCCTGAGGAGAAGTCTGCCGTTACTGCCCTGTGGGGCAAG) ３′ の32Ｐでラベルされた４０塩基対の合成断片で検知し
た。
【００８９】図２は反応Ｉ及びII用の検知されたナイロ
ン膜のオートラジオグラフである。レーン１は０．１ピ
コモルの５８塩基対の対照合成断片で、そのうちの１つ
の鎖は上記プローブと相補的である。レーン２は第１の
増幅サイクルの前の４μｌの反応１の液である。レーン
３は２０回の増幅サイクル後の４μｌの反応１の液であ
る。レーン４は５回の増幅サイクル後の４μｌの反応II
の液である。
【００９０】レーン５はα−32Ｐ−デオキシＮＴＰ及び
ポリメラーゼでラベルされたｐＢＲ３２２（ニューイン
グランドバイオラブ社）のＦｏｋＩ（ニューイングラン
ドバイオラブ社）から成る分子量標準である。レーン３
は、２０サイクル後反応混合物Ｉは適切な分子量を有す
る特定の配列を大量に含み、他の検出できる生成物がな
いことを示している。５サイクル後の反応混合物IIもレ
ーン４に示す通り出発物質である核酸と他の生成物の他
にこの生成物も含んでいる。
【００９１】５サイクル後の反応IIからVIの液５．０μ
ｌに上述の各プライマー５ピコモルを加えた。溶液を４
分間１００℃に加熱し室温へ戻した。それぞれ３ピコモ
ルのα−32Ｐ−デオキシＡＴＰ，α−32Ｐ−デオキシＣ
ＴＰ，α−32ＰデオキシＧＴＰ及びα−32Ｐ−ＴＴＰ、
並びに４単位のクレノー断片を加えた。最終的な容積が
１０μｌであり塩濃度が上記した通りである反応を１０
分間行わせた。ポリメラーゼ活性は６０℃で２０分間加
熱すると失われた。反応II−VIの反応液４μｌを、０．
０８９Ｍトリス硼酸塩緩衝液、２．５ｍＭＥＤＴＡ中で
１２％ポリアクリルアミドゲルに加えた。このゲルを２
５ボルト／cm、４時間電気泳動させ、その後オートラジ
オグラフ処理した。
【００９２】図３は、電気泳動のオートラジオグラフで
ある。レーン１は分子量標準、レーン２は反応II、レー
ン３は反応III 、レーン４は反応IV及びレーン５は反応
Ｖである。対照としてのＤＮＡを伴わない反応VIのレー
ンはレーンのどこにもイメージを有さない。
【００９３】図から、標的ＤＮＡから予想される９４塩
基対断片は、無傷のβ−グロブリンＤＮＡ配列が増幅用
に使用できるときのみ存在できることが分かる（つまり
レーン２のｐＢＲ３２８：ＨｂＡ、レーン３のｐＢＲ３
２８：ＨｂＳ及びレーン５のｐＢＲ３２８：ＨｂＳ／Ｍ
ｓｔII）。ＭｓｔIIによる消化はｐＢＲ３２８：ＨｂＡ
を９４塩基対配列中で切断し、それを増幅できないよう
にし、９４塩基対のバンドはレーン４に現れない。これ
に対し、ｐＢＲ３２８：ＨｂＳの９４塩基対配列はプラ
スミドがＭｓｔIIで消化されても切断せず、従って図１
２に示すように増幅に利用できる。
【００９４】図５〜図１０は９４塩基対配列を増幅する
３サイクルの連鎖反応を示すものである。ＰＣ０１とＰ
Ｃ０２はプライマーＡ及びＢである。右の数はサイクル
を示し、左の数は特定の分子が生産されたサイクル数を
示す。
【００９５】実施例３ 本実施例は、ヒトヘモグロビン遺伝子中の対立遺伝子Ｍ
ｓｔII部位を含む１１０塩基対配列の増幅を示すもので
ある。プライマーは実施例２の技術で調製されたもので
ある。１．０マイクログラムの全ヒトＤＮＡ，１００ピ
コモルのd(ACACAACTGTGTTCACTAGC) 及び１００ピコモル
のd(CAACTTCATCCACGTTCACC) を以下のような１００μｌ
の溶液に溶解させた。
【００９６】１．５ｍＭ各４つのデオキシリボヌクレオ
シド三リン酸 ３０ｍＭpH７．９のトリスアセテート緩衝液 ６０ｍＭ酢酸ナトリウム １０ｍＭ酢酸マグネシウム ２５ｍＭジチオスレイトール
【００９７】この溶液を１００℃で１分間加熱し、迅速
に２５℃に下げて１分間加熱し、その後ＤＮＡポリメラ
ーゼのクレノー断片２．５単位を加えた。ポリメラーゼ
の反応が２５℃で２分間行い、その後加熱、冷却、クレ
ノー断片の添加及び反応を望むだけ繰り返した。各サイ
クルの効率が７０℃で、１５サイクル行って、β−グロ
ビン遺伝子の所望の１１０塩基対断片１．４フェトモル
を合成した。
【００９８】実施例４ 本実施例は、ヒトヘモグロビン遺伝子の対立遺伝子中の
ＭｓｔII部位を含む２４０塩基対配列の増幅を示すもの
である。この配列は、ＮｃｏＩ，ＨｉｎｆＩ及びＭｓｔ
II制限部位を含んでいる。pHが８．０で、６０ｍＭ酢酸
ナトリウム、３０ｍＭトリスアセテート及び１０ｍＭ酢
酸マグネシウムの混合物（０．１ピコモルのｐＢＲ３２
８：ＨｂＡを含む）に、１００ピコモルのd(GGTTGGCCAA
TCTACTCCCAGG) プライマー、１００ピコモルのd(TAACCT
TGATACCAACCTGCCC) プライマー、各１０００ピコモルの
デオキシＡＴＰ、デオキシＣＴＰ、デオキシＧＴＰ及び
ＴＴＰを含む２μｌの溶液Ａを加えた。
【００９９】２つのプライマーは実施例２に記載した技
術で調製した。溶液を１００℃で４分間加熱し、空気中
で２分間冷却し、その後Ｅ．コーリーＤＮＡポリメラー
ゼのクレノー断片４単位を含む液１μｌを加えた。反応
を１０分間進行させ、その後溶液Ａの添加、加熱、冷
却、ポリメラーゼの添加及び反応からなるサイクルを３
回繰り返した。反応液５．０μｌに、上記の各オリゴヌ
クレオチドプライマー５ピコモルを加えた。溶液を１０
０℃で４分間加熱し、室温まで下げ、その後それぞれ３
ピコモルのα−32Ｐ−ラベルされたデオキシリボヌクレ
オシド三リン酸及び４単位のクレノー断片を加えた。最
終的な容量が１０μｌで塩濃度が上記の通りである反応
を１０分間進行させる。
【０１００】ポリメラーゼ活性は６０℃で２０分間加熱
すると失活した。２μｌのアリコートをＮｃｏＩ、Ｈｉ
ｎｆＩ及びＭｓｔIIで消化し、pH８．３の０．０８９Ｍ
トリスアセテート緩衝液、０．２５ｍＭＥＤＴＡ中で１
２％ポリアクリルアミドゲルに加えた。ゲルを２５ボル
ト／cmで４時間電気泳動させ、オートラジオグラフ処理
した。図１２は電気泳動のオートラジオグラフを示し、
ここでレーン１は分子量標準、レーン２は酵素の消化を
伴わないもの（無傷の２４０塩基対）、レーン３はＮｃ
ｏＩによる消化（１３１及び１０９塩基対）、レーン４
はＭｓｔIIによる消化（１４９及び９１塩基対）、そし
てレーン５はＨｉｎｆＩによる消化（１４４及び９６塩
基対）である。オートラジオグラフは２４０塩基対反応
の増幅したものと一致する。
【０１０１】実施例５ 本実施例は、逐次的消化による鎌状赤血球貧血を検出す
るための本発明の方法の使用を示すものである。オリゴデオキシリボヌクレオチドの合成及びリン酸化 ５′* CTGACTCCTGAGGAGAAGTCTGCCGTTACTGCCCTGTGGG ３′ の配列のラベルされたＤＮＡプローブ（* がラベルを意
味する）ＲＳ０６、及びＲＳ０６と３つの塩基対がミス
マッチしている ３′GACAGAGGTCACCTCTTCAGACGGCAATGACGGGACACCC ５′ の配列のラベルされていないブロックオリゴマーＲＳ１
０を、実施例２（Ｉ）の方法に従って合成した。
【０１０２】プローブＲＳ０６は、その５ピコモルを、
７０ｍＭトリス緩衝液（pH７．６），１０ｍＭＭｇＣｌ
2 ，１．５ｍＭスペルミン及び２．５ｍＭジチオスレイ
トールを含む反応容量４０μｌ中の４単位のＴ４ポリヌ
クレオチドキナーゼ（ニューイングランドバイオラブ
社）及び５０ピコモルのγ−32Ｐ−ＡＴＰ（ニューイン
グランドニュークレア社、約７２００Ｃｉ／ミリモル）
と接触させることによりラベルした。
【０１０３】全容量を２５ｍＭＥＤＴＡで１００μｌに
調節し、トリス−ＥＤＴＡ（ＴＥ）緩衝液（１０ｍＭト
リス緩衝液、０．１ｍＭＥＤＴＡ，pH８．０）により平
衡化されたバイオラッド製の１mlのＢｉｏ Ｇｅｌ Ｐ
−４スピン透析カラム上でマニアティスらがＭｏｌｅｃ
ｕｌａｒ Ｃｌｏｎｉｎｇ４６４−４６５頁（１９８２
年）に記載している方法に従って精製した。ラベルされ
たプローブは、トリス−硼酸−ＥＤＴＡ（ＴＢＥ）緩衝
液（８９ｍＭトリス、８９ｍＭ硼酸、２．５ｍＭＥＤＴ
Ａ，pH８．３）中１８％のポリアクリルアミドゲル（１
９：１のアクリルアミド：ＢＩＳとバイオラッド）上で
５００ｖｈｒにて電気泳動してさらに精製した。
【０１０４】オートラジオグラフによる位置きめの後、
ラベルされたプローブを含む部分を切り取り、粉砕し、
０．２mlのＴＥ緩衝液中へ一晩かけて４℃で溶出させ
た。反応生成物のＴＣＡ沈澱は比活性が４．９Ｃｉ／ミ
リモルであり、最終的濃度が２０ピコモル／mlであるこ
とを示している。ラベルされないＲＳ１０ブロッキング
オリゴマーは２００ピコモル／mlの濃度で使用した。
【０１０５】細胞系からのヒトゲノムＤＮＡの分離 実質的にステットラーらのＰＮＡＳ７９巻５９６６−５
９７０頁（１９８２年、Ｍｏｌｔ４について）に記載の
方法及びマニアティスらのＭｏｌｅｃｕｌａｒＣｌｏｎ
ｉｎｇ２８０−２８１頁（１９８２年）に記載の方法を
使用して、Ｍｏｌｔ４、ＳＣ−１及びＧＭ２０６４のリ
ンパ球系から高分子のゲノムＤＮＡを分離した。
【０１０６】Ｍｏｌｔ４（ヒューマン・ミュータント・
セル・デポジトリー、ＧＭ２２１９Ｃ）は正常のβ−グ
ロビンについてホモ接合体のＴ細胞系であり、そしてＡ
ＴＣＣに１９８５年３月１９日に寄託されたＳＣ−１は
鎌状赤血球貧血対立遺伝子についてホモ接合体のＥＢＶ
で形質変換されたＢ細胞系である。ＧＭ２０６４（ヒュ
ーマン・ミュータント・セル・デポジトリー、ＧＭ２０
６４）は胎児ヘモグロビンの遺伝的な永続性（ＨＰＦ
Ｈ）についてホモ接合体である個体当初単離され、β−
又はδ−グロビン遺伝子配列を含んでいない。全ての細
胞系は１０％の牛胎児血清を含むＲＰＭＩ−１６４０中
に維持された。
【０１０７】臨床血液試料からのヒトゲノムＤＮＡの単
離 既知の鎌状細胞キャリヤー（ＡＳ）からのＣＨ１２と名
付けられた臨床血液試料をカルフォルニア州オークラン
ドの小児病院のベルトラム・ルビン博士から得た。ヌン
ベルグらのＰｒｏｃ．Ｎａｔ．Ａｃａｄ．Ｓｃｉ．７５
巻５５５３−５５５６頁（１９７８年）に記載されてい
る方法の変法を使用して、主に末梢の血液リンパ球から
成るバフィーコート部分からゲノムＤＮＡを調製した。
【０１０８】細胞を、５mlのトリス−ＥＤＴＡ−ＮａＣ
ｌ（ＴＥＮ）緩衝液（pH８の１０ｍＭトリス、pH８，１
ｍＭＥＤＴＡ、１０ｍＭＮａＣｌ）中に再懸濁し、０．
２mg／mlのプロテイナーゼ、０．５％のＳＤＳに調節
し、そして３７℃で一晩インキュベートした。過塩素酸
ナトリウムを０．７Ｍに加え、そして細胞溶解物を室温
で１−２時間穏やかに振とうした。
【０１０９】細胞溶解物をフェノールとクロロフォルム
の１：１混合物３０mlで抽出し、続いてクロロフォルム
３０mlで抽出し、次にエタノールで核酸を沈澱させた。
ペレットを２mlのＴＥ緩衝液に再懸濁させ、ＲＮａｓｅ
を０．００５mg／mlに加えた。３７℃で１時間消化させ
た後、ＤＮＡを同量のフェノール、フェノール／クロロ
フォルム、及びクロロフォルムでそれぞれ一度ずつ抽出
し、エタノールで沈澱させた。ＤＮＡを０．５mlのＴＥ
緩衝液に再懸濁させ、２６０nmの吸収により濃度を決定
した。
【０１１０】β−グロビン配列を選択的に増幅するため
のポリメラーゼ連鎖反応 ２マイクログラムのゲノムＤＮＡを、１０ｍＭトリス緩
衝剤（pH７．５），５０ｍＭＮａＣｌ，１０ｍＭＭｇＣ
ｌ2 ，１５０ピコモルの配列d(CACAGGGCACTAACG)のプラ
イマーＡ、及び配列d(CTTTGCTTCTGACACA) のプライマー
Ｂを含む反応容量１００μｌの当初溶液中で増幅し、か
つ蒸発を防ぐため約１００μｌ厚の鉱油で被覆した。各
ＤＮＡ試料につき、１サイクルが次の３ステップから成
る増幅のための１５サイクルを行った。
【０１１１】（１）２分間９５℃で熱ブロックセット中
で変性する。 （２）熱ブロックセットを直ちに３０℃に移し２分間プ
ライマーとゲノムＤＮＡがアニーリングするようにす
る。 （３）Ｅ．コーリーＤＮＡポリメラーゼＩのクレノー断
片（ニューイングランドバイオラブ）５単位とデオキシ
ＡＴＰ、デオキシＣＴＰ、デオキシＧＴＰ及びＴＴＰそ
れぞれ１ナノモルを含む２μｌの溶液（１０ｍＭトリス
（pH７．５）、５０ｍＭＮａＣｌ、１０ｍＭＭｇＣ
ｌ2 、及び４ｍＭジチオスレイトールから成る緩衝液
中）を加える。この伸長反応を３０℃にて１０分間行っ
た。最後のサイクルの後、９５℃に２分間維持して反応
を停止させた。鉱油は０．２mlのクロロフォルムで抽出
して廃棄した。最後の反応液の容量は１３０μｌであっ
た。
【０１１２】プローブ及びＤｄｅＩ／ＨｉｎｆＩによる
増幅したゲノムＤＮＡのハイブリダイゼーション／消化 ２５マイクロリットルの増幅されたゲノムＤＮＡをエタ
ノールで沈澱させ、同量のＴＥ緩衝液中に再懸濁した。
１０マイクロリットル（１５４ngのゲノムＤＮＡと同等
の前増幅体を含む）を１．５mlのマイクロフュージ管に
入れ、そして２０μｌのＴＥ緩衝液により最後の容量を
３０μｌとした。試料を鉱油で被覆して９５℃で１０分
間変性した。ラベルされたＲＳ０６プローブ０．０２ピ
コモルを含む０．６ＭＮａＣｌ１０マイクロリットルを
管に加え、穏やかに混合し、直ちに５６℃の熱ブロック
に移して１時間おいた。
【０１１３】ラベルしていないＲＳ１０ブロッキングオ
リゴマー４マイクロリットル（０．８ピコモル）を加
え、更に１０分間同じ温度でハイブリダイゼーションを
続けた。５マイクロリットルの６０ｍＭＭｇＣｌ2 ／
０．１％ＢＳＡ及び１μｌのＤｅｌＩ（１０単位、ニュ
ーイングランドバイオラブ）を加え、再アニーリングさ
れたＤＮＡを５６℃で３０分間消化した。１マイクロリ
ットルのＨｉｎｆＩ（１０単位、ニューイングランドバ
イオラブ）を加え、更に３０分インキュベートした。４
μｌの７５ｍＭＥＤＴＡと６μｌのトラッキング染料を
最終容積が６１μｌになるように反応混合物に加えて反
応を終了した。
【０１１４】鉱油を０．２mlのクロロフォルムで抽出
し、１８μｌの反応混合物（４５nmのゲノムＤＮＡ）を
ヘーファーＳＥ２００装置中の３０％ポリアクリルアミ
ドのミニゲル（１９：１、バイオラド）に負荷した。こ
のゲルをブロモフェノールブルー染料の前端が当初の位
置から３．０cm動くまで約３００ボルトで１時間電気泳
動させた。該ゲルの前端の１．５cmは取り除かれ、残り
のゲルは４日間−７０℃で強化スクリーンに曝される。
【０１１５】写真の検討（図１６） 各レーンは４５ngの増幅されたゲノムＤＮＡを含んでい
る。レーンＡはＭｏｌｔ４ＤＮＡを、レーンＢはＣＨ１
２を、レーンＣはＳＣ−１を、又レーンＤはＧＭ２０６
４を含んでいる。Ｍｏｌｔ４は、細胞当たり２コピーの
βA 遺伝子を有する正常の個体の遺伝子型ＣＡＡであ
り、ＣＨ１２は、細胞当たり１個のβA と１個のβS 遺
伝子を有する鎌状細胞キャリアからの臨床用試料（Ａ
Ｓ）であり、そしてＳＣ−１は細胞当たり２コピーのβ
S を有する鎌状血球血貧血個体の遺伝子型を意味する。
ＧＭ２０６４はβ−又はδ−グロビン配列を含有せず、
ネガティブ対照として存在する。
【０１１６】写真から分かるようにＤｄｅＩで開裂され
た、βA 特異的であるオクタマーはβA 遺伝子を含むＤ
ＮＡにのみ存在し（レーンＡ及びＢ）、ＨｉｎｆＩで開
裂された、βS 特異性を有するトリマーは、βS 遺伝子
を含むＤＮＡにのみ存在する（レーンＢ及びＣ）。トリ
マー及びオクタマーの両者の存在（レーンＢ）は鎌状赤
血球貧血キャリアを示すものであり、オクタマーのみを
生ずる正常の個体（レーンＡ）及びトリマーのみを示す
鎌状赤血球貧血にかかっている個体（レーンＣ）から区
別される。比較のため、上記実験を増幅されていないゲ
ノムＤＮＡを用いて繰り返し行い、増幅を行うと検出感
度が少なくとも１０００倍増加することが分かった。
【０１１７】実施例６ 本実施例は、ラベルされたプローブを使用することなく
全ヒトＤＮＡ中の全く精製されていない単一コピー遺伝
子をゲル上で直接検出する方法を示すものである。実施
例３に記載した技術を用い、β−グロブリン遺伝子の第
１エクソン中の配列からの１１０塩基対断片を、全ヒト
ＤＮＡ１０マイクログラムから２０サイクルで増幅し
た。この２０サイクル後に生産される１１０塩基対断片
は、臭化エチジウムにより容易に染色されてゲル上で見
ることができた。配列は、最初に制限酵素ＤｄｅＩによ
り切断されると、配列がβ−グロビンのＳ対立遺伝子中
における場合のように酵素により認識される制限部位を
含まないものでない限り、増幅されなかった。
【０１１８】実施例７ Ａ．ヒトβ−グロビンＡ対立遺伝子からの１．９kb挿入
部を含有する合計１００フェムトモルのｐＢＲ３２８，
５００Ｃｉ／モルである各α−32Ｐ−デオキシＮＴＰを
５０ナノモルずつ、及び実施例３で使用した各プライマ
ー１ナノモルを、１００μｌの３０ｍＭトリス−アセテ
ート（pH７．９）、６０ｍＭ酢酸ナトリウム、１００ｍ
Ｍジチオスレイトール及び１０ｍＭ酢酸マグネシウムを
含む溶液中に溶かした。
【０１１９】この溶液を１００℃にして２分加熱し、２
５℃にて１分冷却した。４．５単位のＥ．コーリーＤＮ
ＡポリメラーゼＩ及び０．０９単位の無機ピロフォスフ
ァターゼを加えて反応混合物中でピロリン酸が生ずるの
を防止し、その後反応を２５℃で２分間進行させ、更に
加熱、冷却、酵素の添加及び反応のサイクルを９回繰り
返した。
【０１２０】各合成サイクルの後、１０μｌのアリコー
トを取り出し１μｌの６００ｍＭＥＤＴＡに加えた。そ
れぞれを、９０ｍＭのトリスボレート及び２．５ｍＭＥ
ＤＴＡ中、pH８．３で１４％のポリアクリルアミドゲル
上で２４ボルト／cm、２．５時間で分析した。操作の終
了したゲルは、０．５μｇ／mlの臭化エチジウムを加え
た同じ緩衝液に２０分浸し、当初の緩衝液で洗浄し、赤
フィルターを用いて紫外線中で写真を撮影した。
【０１２１】生産された１１０塩基対断片は紫外線でゲ
ルから切り出し、そしてクレンコフ放射により計数し
た。Ｎがサイクル数を意味し、ｙがサイクル毎の部分的
収率である式 ｐｍｏｌｅｓ／１０μｌ＝０．０１（（１＋ｙ）N −ｙ
Ｎ−１）、 にデータを一致させようとする試みは、ｙが０．６１９
であるときに楽観的なものとなる。これは、十分な増幅
が起こっていることを暗示している。
【０１２２】Ｂ．各デオキシＮＴＰを１００ナノモルず
つ１００μｌの反応溶液に加え、放射性ラベルを行わ
ず、各サイクル毎に液を取り出さなかった以外は、上記
実験を繰り返した。１０サイクル後に反応物を２分間沸
騰させて反応を停止させ、５７℃，１時間で再ハイブリ
ダイゼーションを行った。１１０塩基対生成物の配列
を、その８μｌのアリコートを、１μｌのウシ血清アル
ブミン（２５mg／ml）と１μｌの好適な制限酵素（Ｈｉ
ｎｆＩ，ＭｎｌＩ，ＭｓｔII，ＮｃｏＩ）を加えて制限
分析し、３７℃で１５時間反応させて確認した。ＰＡＧ
Ｅは、上述の通り行った。
【０１２３】実施例８ 本実施例は、ｐＢＲ３２８とｐＢＲ３２２の種々の断片
を増幅するために異なったプライマーを使用する例を示
す。Ａ．次のプライマーを使いｐＢＲ３２８の１３０塩
基対断片を調製すること以外は実施例７Ａと同じように
実験を繰り返した。 d(TTTGCTTCTGACACAACTGTGTTCACTAGC) 及び d(GCCTCACCACCAACTTCATCCACGTTCACC)
【０１２４】Ｂ．次のプライマーを用いたこと以外は実
施例７Ａと同じように実験を繰り返しｐＢＲ３２８の２
６２塩基対断片を調製した。反応時間はサイクル当たり
２０分であった。 d(GGTTGGCCAATCTACTCCCAGG) 及び d(TGGTCTCCTTAAACCTGTCTTG)
【０１２５】Ｃ．ヒトβ−グロビンＳ対立遺伝子からの
１．９kbの挿入部を含む１００フェムトモルのｐＢＲ３
２８のＭｓｔII消化物を当初の鋳型として用いた以外
は、実施例８Ｂと同様に実験を行った。該プラスミドは
ＭｓｔIIにより数回切断されたが、増幅すべき配列の内
側では切断が起こらなかった。更に、使用したプライマ
ーは次の通りで、２４０塩基対断片を生産した。 d(GGTTGGCCAATCTACTCCCAGG) 及び d(TAACCTTGATACCAACCTGCCC)
【０１２６】Ｄ．１００フェムトモルのｐＢＲ３２２の
ＮｒｕＩ消化物を鋳型として用い、１００μｌの反応液
中で各デオキシＮＴＰを２００ナノモル使用し、次のプ
ライマーを使用してｐＢＲ３２２から５００塩基対断片
を生産した以外は実施例７Ｂと同様に実験を行った。 d(TAGGCGTATCACGAGGCCCT) 及び d(CTTCCCCATCGGTGATGTCG)反応時間は３７℃でサイクル
当たり２０分であった。最後の再ハイブリダイゼーショ
ンは５７℃で１５時間行った。電気泳動は４％アガロー
スゲル上で行った。
【０１２７】実施例９ 本実施例は、インビトロ変異が増幅されたセグメントに
導入されるような本発明方法を例示するものである。 Ａ．ＮｒｕＩで直線化したｐＢＲ３２２合計１００フェ
ムトモル、１ナノモルの７５塩基対断片を生成するよう
に設計されたそれぞれ次式 d(CGCATTAAAGCTTATCGATG) 及び d(TAGGCGTATCACGAGGCCCT) のプライマー、それぞれ１００ナノモルの各デオキシＮ
ＴＰを、pH８の４０ｍＭトリス、２０ｍＭＭｇＣｌ2 ，
５ｍＭジチオスレイトール及び５mg／mlのウシ血清アル
ブミンの溶液１００μｌ中で混合した。
【０１２８】この混合物を１００℃にして１分間加熱
し、水浴中２３℃，０．５分間冷却し、次に４．５単位
のクレノー断片と０．０９単位の無機ピロフォスファタ
ーゼを加え、反応を３分間行った。加熱、冷却、酵素添
加及び反応のサイクルを９回繰り返した。１０回目の反
応サイクルは凍結により終了させ、反応混合物のアリコ
ート８μｌを４％アガロースゲルに適用し、臭化エチジ
ウムにより視覚化した。
【０１２９】Ｂ．オリゴヌクレオチドプライマーとして
次式のものを使用した以外は実施例９Ａと同様の実験を
繰り返した。 d(CGCATTAAAGCTTATCGATG) 及び d(AATTAATACGACTCACTATAGGGAGATAGGCGTATCACGAGGCCCT) これらのプライマーは１０１塩基対を生産するように設
計され、その（２番目のプライマー中の）２６ヌクレオ
チドはｐＢＲ３２２には存在しない。
【０１３０】これらのヌクレオチドはＴ７プロモーター
の配列を表すもので、これを、ｐＢＲ３２２からの７５
塩基対配列に、２０の相補的塩基と２６塩基の５′側伸
長部とを有するプライマーを使用して連結した。この方
法は２時間より少ない時間で実施することができ、１０
０フェムトモルのｐＢＲ３２２から比較的純粋な１０１
塩基対断片２ピコモルを生産することができた。Ｔ７プ
ロモーターはＲＮＡ転写を開始させるために使用でき
る。Ｔ７ポリメラーゼを１０１塩基対断片に加えて単鎖
ＲＮＡを生成せしめることができる。
【０１３１】Ｃ．オリゴヌクレオチドプライマーとして
下記のものを使用して、ｐＢＲ３２２から１０００塩基
対断片を調製した以外は実施例８Ｄと同様に実験を繰り
返した。 d(TAGGCGTATCACGAGGCCCT) 及び d(CCAGCAAGACGTAGCCCAGC)
【０１３２】Ｄ．上記９Ｃと同様の実験を繰り返した。
但し、オリゴヌクレオチドプライマーとして下記のも
の、 d(TAGGCGTATCACGAGGCCCT) 及び d(AATTAATACGACTCACTATAGGGAGATAGGCGTATCACGAGGCCCT) を使用して１０２６対断片を調製した。２番目のプライ
マーの２６ヌクレオチドはｐＢＲ３２２には存在せず、
上記のＴ７プロモーターを示すものである。このプロモ
ーターは、ｐＢＲ３２２からの１０００塩基対断片に隣
接して挿入された。
【０１３３】これらの結果は、鋳型鎖と完全にマッチし
ていないがそれにもかかわらず十分にハイブリダイズし
て酵素的に伸長するプライマーは、当初の鋳型に対応す
る鎖よりむしろプライマーの鎖を含む長鎖生成物を生成
せしめるということを暗示する。長鎖生成物はインビト
ロ変異を生じさせる第２のプライマー用の鋳型としての
役割を果たす。その後のサイクルでは、更に多くのミス
ペアしたプライミングが要求されないので、効率が減少
することなくこの変異は増幅される。この場合、その
５′末端に相補的でない伸長部分があるプライマーが、
複製されるべき鋳型に隣接して生成物中に新しい配列を
挿入するために使用された。
【０１３４】実施例１０ 本実施例は単コピー遺伝子を増幅させる際にバックグラ
ウンドを減少させるためにネスト状に（ｎｅｓｔｅｄ）
セットしたプライマーを使用することを例示するもので
ある。野性型β−グロビン対立遺伝子についてホモ接合
体である全ヒトＤＮＡに対して、２０サイクルの増幅を
次のように行った。
【０１３５】１０μｇのＤＮＡ、それぞれ２００ピコモ
ルの次式のプライマー、 d(ACACAACTGTGTTCACTAGC) 及び d(CAACTTCATCCACGTTCACC) 並びに１００ナノモルずつのｄＮＴＰを、１００μｌの
３０ｍＭトリス−アセテート、６０ｍＭ酢酸ナトリウ
ム、１０ｍＭジチオスレイトール、及び１０ｍＭ酢酸マ
グネシウム中で１００℃にて１分間加熱し、２５℃に１
分間下げて、そして２単位のクレノー断片とともに２分
間処理した。加熱、冷却、クレノー試薬の添加のサイク
ルを１９回繰り返した。
【０１３６】１０μｌの液体を反応混合物から取り出
し、更に１０回の増幅のためのサイクルを次の各プライ
マーを用いて行った。 d(CAGACACCATGGTGCACCTGACTCCTG)及び d(CCCCACAGGGCAGTAACGGCAGACTTCTCC) これらは、上記で生産された１１０塩基対断片中に含ま
れる５８塩基対断片を増幅した。増幅すべき最後の１０
回のサイクルは、１０μｌのアリコートを、上記した各
デオキシＮＴＰ １００ナノモルと各プライマー２００
ピコモルを含む９０μｌの新しいトリス−アセテート緩
衝液に希釈することにより達成することができた。
【０１３７】反応条件は上記の通りとした。１０サイク
ルの後１０μｌのアリコート（当初のＤＮＡの１００ナ
ノグラムに対応）を６％のＮｕシーブ（ＦＭＣ社）アガ
ロースゲルに加え、臭化エチジウムを使って視覚化し
た。図１７は、紫外線で発光させた従来法の通り赤いフ
ィルターを通して写真撮影した上記ゲルを示すものであ
る。レーン１は分子量のマーカーである。レーン２は上
記反応のアリコートである。レーン３は当初の野性型Ｄ
ＮＡが増幅の前にＤｄｅＩにより開裂されたこと以外は
上記したものと同じ反応のアリコートである。
【０１３８】レーン４は鎌状赤血球貧血β−グロビン対
立遺伝子についてホモ接合体であるヒトＤＮＡを増幅の
前にＤｄｅＩで処理したこと以外は上記と同様な反応の
アリコートである（鎌状赤血球貧血対立遺伝子は増幅さ
れる断片内にＤｄｅＩ部位を含まない）。レーン５は鮭
の精子ＤＮＡでヒトＤＮＡを置き換えた以外は上記と同
様の反応のアリコートである。レーン６は増幅後反応液
をＤｄｅＩで処理したこと以外は上記と同様な反応のア
リコートである（ＤｄｅＩは５８塩基対の野性型生成物
を２７塩基対及び３４塩基対の断片に変換する）。レー
ン７は増幅後ＤｄｅＩで処理したレーン４の材料のアリ
コートである（５８塩基対の鎌状赤血球貧血生成物はＤ
ｄｅＩを含まない）。
【０１３９】アガラーセゲルの臭化エチジウム染色のみ
を使用してヒトＤＮＡの１マイクログラムからの単コピ
ー遺伝子を代表する５８塩基対断片を検出するために
は、約５００，０００倍に増幅することが必要である。
これは、ここで２つのオリゴヌクレオチドのネスト状セ
ットを使用して達成することができる。第１のセットは
１１０塩基対断片を増幅し、そして内部のネスト状セッ
トは、図１７に示すように便利に検出できるレベルにな
るまでこの生成物のサブ−断片を増幅する。
【０１４０】先行する増幅工程で増幅された配列中に含
まれ、又他のプライマーの伸長生成物中にも含まれるよ
り小さな配列をプライマーを使って増幅する本法は、例
えばコナーらのＰＮＡＳ８０巻２７８頁（１９８３年）
及びレアリーらのＰＮＡＳ８０巻４０４５頁（１９８３
年）に記載されているように放射性同位体又は非放射性
同位体プローブのハイブリダイゼーションの方法論に頼
ることなく、β−グロビンの座における野性型を鎌状赤
血球貧血対立遺伝子から区別することを可能にする。
【０１４１】実施例１１ 本法は、患者のＤＮＡ試料中の例えばクラミジアのよう
な伝染性疾患と関連する特定の配列を、所望の増幅され
た配列を含むビオチン化されたハイブリダイゼーション
プローブを使用しかつ前述の米国特許第４，３５８，５
３５号に記載された方法を使用して検出する際に有用で
あることが期待される。ビオチン化されたハイブリダイ
ゼーションプローブは、一部が二重鎖となったＤＮＡ
に、次式のスペーサーアームを介してビオチンに結合し
た４′−メチレン置換−４，５′−８−トリメチルプソ
ラレンを挿入しかつ光を照射することにより調製するこ
とができる。
【０１４２】
【化１】 式中、Ｒは−Ｈ又はＣＨＯ基であり、Ｒ″は−Ｈであ
り、ｘは１〜４の数であり、そしてｙは２〜４の数であ
る。プローブ上のビオチニル基の検出には、エンゾバイ
オケム社により市販されているストレプタビジン−酸性
フォスファターゼ複合体を用いて、パンフレットに製造
者が示している検出方法により達成することができる。
ハイブリダイゼーションプローブは、検出用複合体との
結合、及びそれに続く酸性ホスファターゼにより触媒さ
れる反応（この反応が沈澱性色素を生成する）に基く沈
澱した染色スポットとして見ることができる。
【０１４３】材料の寄託 細胞系ＳＣ−１（ＣＴＣＣ＃００８２）は、１９８５年
３月１９日、米国、２０８５２、メリーランド州ロック
ビルパークローンドライブ１２３０１に所在するアメリ
カン・タイプ・カルチャー・コレクション（ＡＴＣＣ）
にＡＴＴＴ受理番号第ＣＲＬ８７５６号として寄託され
た。ＳＣ−１の寄託は、ＡＴＣＣと本特許出願人のシー
タス・コーポレーションとの間の契約に従って行われ
た。
【０１４４】ＡＴＣＣとの契約は、本寄託を記載しかつ
特定する米国特許が発行された場合又は米国又は外国特
許出願が公衆に公告された場合又は公開された場合のい
ずれか早い方が来たときにこの細胞系の子孫を公衆がそ
れを永続的に利用できるようにするために提供し、更に
本細胞系を利用させることについては、米国特許商標局
長官が米国特許法第１２２条及びそれに関する長官のル
ール（３７ ＣＦＲ１，１４条も特に８８６ ＯＧ ６
３８に関連して含む）に従って権限を持って決定した人
間に対しても行う。
【０１４５】本出願の譲受人はもし寄託した細胞系が好
適な条件下で培養したにもかかわらず、死滅し、失わ
れ、損傷したときは通知を受けてから迅速に同じ細胞系
の育成培養基と置き換えることに同意する。纏めると、
本発明はまず１つ又はそれ以上の特定の核酸を、プライ
マーの伸長により生産される生成物が引き続き次のプラ
イマーの伸長反応の鋳型としての役割を果たすような連
鎖反応を用いて増幅させることにより核酸中の配列を検
出するようにした方法を提供する。本法は当初にほんの
僅かの量しか含まれていない核酸配列を検出するために
特に有用である。更に増幅法は分子クローニングにも使
用することができる。
【図面の簡単な説明】
【図１】図１は、増幅されることが望まれるヒト−β−
グロビンの９４塩基対長の配列を示すものであり、鎌状
赤血球貧血に伴う単一塩基対変化を９４ｍｅｒの下方に
描いてある。
【図２】図２は、ヒトの野性型ＤＮＡ中、及び正常のβ
−グロビン遺伝子の１．９kbのＢａｍＨＩ断片を含むプ
ラスミド（ｐＢＲ３２８：ＨｂＡと示される）中に含ま
れる上記９４ｍｅｒの増幅を示す臭化エチジウムで染色
されたポリアクリルアミドのゲルの写真であり、電気泳
動図である。
【図３】図３は、ｐＢＲ３２８：ＨｂＡ、及びβ−グロ
ビンの鎌状赤血球貧血対立遺伝子の１．９kbのＢａｍＨ
Ｉ断片を含有するプラスミド（ｐＢＲ３２８：ＨｂＳと
称する）中に存在する特定の標的９４ｍｅｒ配列のいず
れかの増幅を示すポリアクリルアミドゲル電気泳動図の
オートラジオグラフを示し、ｐＢＲ３２８：ＨｂＡでは
増幅されるべき配列がＭｓｔIIにより開裂され、そして
ｐＢＲ３２８：ＨｂＳでは増幅されるべき配列が処理さ
れたがＭｓｔIIにより開裂されなかった。
【図４】図４は図５〜図１０の相互位置関係を示す図で
ある。
【図５】図５は、２つのオリゴヌクレオチドプライマー
を用いる３サイクルについて、ヒトβ−グロビンの所望
の９４ｍｅｒ配列の増幅のためのポリメラーゼの連鎖反
応のステップと生成物の詳細を示すものである。
【図６】図６は、２つのオリゴヌクレオチドプライマー
を用いる３サイクルについて、ヒトβ−グロビンの所望
の９４ｍｅｒ配列の増幅のためのポリメラーゼの連鎖反
応のステップと生成物の詳細を示すものである。
【図７】図７は、２つのオリゴヌクレオチドプライマー
を用いる３サイクルについて、ヒトβ−グロビンの所望
の９４ｍｅｒ配列の増幅のためのポリメラーゼの連鎖反
応のステップと生成物の詳細を示すものである。
【図８】図８は、２つのオリゴヌクレオチドプライマー
を用いる３サイクルについて、ヒトβ−グロビンの所望
の９４ｍｅｒ配列の増幅のためのポリメラーゼの連鎖反
応のステップと生成物の詳細を示すものである。
【図９】図９は、２つのオリゴヌクレオチドプライマー
を用いる３サイクルについて、ヒトβ−グロビンの所望
の９４ｍｅｒ配列の増幅のためのポリメラーゼの連鎖反
応のステップと生成物の詳細を示すものである。
【図１０】図１０は、２つのオリゴヌクレオチドプライ
マーを用いる３サイクルについて、ヒトβ−グロビンの
所望の９４ｍｅｒ配列の増幅のためのポリメラーゼの連
鎖反応のステップと生成物の詳細を示すものである。
【図１１】図１１は図５〜図１０に示す増幅におけるサ
イクル数とＤＮＡコピー数との関係を示す。
【図１２】図１２は、ｐＢＲ３２８：ＨｂＡ中の２４０
ｍｅｒ配列の４サイクル後の増幅を示す臭化エチジウム
で染色されたポリアクリルアミドのゲルを示す電気泳動
図の写真であり、ここはアリコートがＮｃｏＩ（レーン
３）、ＭｓｔII（レーン４）又はＨｉｎｆＩ（レーン
５）により消化される。レーン１は分子量の基準で、レ
ーン２は無傷の２４０ｂｐの生成物を含んでいる。
【図１３】図１３は、ＤｄｅＩ及びＨｉｎｆＩ制限部位
間にある正常な（βA ）β−グロビン遺伝子及び鎌状赤
血球（βS ）β−グロビン遺伝子の配列を示すもので、
βA についての１本線はＤｄｅＩ部位（CTGAG)の位置を
示し、βA 及びβS についての２重線はＨｉｎｆＩ部位
（GACTC)の位置を示している。
【図１４】図１４は、４０ｍｅｒプローブ、並びにＤｄ
ｅＩ及びこれに続くＨｉｎｆＩ制限酵素を用いる正常β
−グロビンの逐次的な消化の結果を示すものである。
【図１５】図１５は、図１４と同じ４０ｍｅｒプローブ
並びにＤｄｅＩ及びこれに続くＨｉｎｆＩ制限酵素を使
用する鎌状β−グロビンの逐次的な消化の結果を示すも
のである。
【図１６】図１６は、この発明の増幅にかけられた全ヒ
トＤＮＡの試料中に存在するβ−グロビン対立遺伝子を
特異的に特徴付けるための、図１４と同じ４０ｍｅｒプ
ローブの使用を示す、臭化エチジウムで染色されたポリ
アクリルアミドのゲルを示す電気泳動図の写真である。
【図１７】図１７は、臭化エチジウムと紫外線を用いて
視覚化した６％のＮｕシーブアガロースゲルの電気泳動
図の写真を示すものである。この写真は、１１０−ｂｐ
増幅生成物のサブ−フラグメントの増幅を示し、このサ
ブ−フラグメントは１１０ｂｐ断片内の内部ネストであ
る。
───────────────────────────────────────────────────── フロントページの続き (56)参考文献 ＮＵＣＬＥＩＣ ＡＣＩＤＳ ＲＥＳ ＥＡＲＣＨ ＶＯＬ．10，ＮＯ．３，Ｐ Ｐ1029−1038（1982)

Claims (13)

    (57)【特許請求の範囲】
  1. 【請求項１】 核酸又は核酸混合物中の少なくとも１つ
    の特定の核酸を増幅するためのキットであって、ここで
    各核酸は同一の長さもしくは異る長さの２つの分離され
    た相補的な鎖又は１本鎖から成り、該キットは少なくと
    も１対のオリゴヌクレオチドプライマーを含んで成り、
    ここで、 ａ）前記プライマー対の一方のプライマーは前記１本鎖
    核酸に対してもしくは前記２本鎖核酸の一方の鎖に対し
    て実質的に相補的であり、 ｂ）前記プライマー対の他方のプライマーは前記１本鎖
    核酸の相補体に対して又は前記２本鎖核酸の他方の鎖に
    対して実質的に相補的であり、 ｃ）前記プライマーの少なくとも一方は、 （ａ）その５′−末端に結合したヌクレオチド配列をさ
    らに含有し、該配列は前記増幅されるべき核酸配列に対
    して非−相補的であり、又は （ｂ）塩基又はヌクレオチド配列により中断されてお
    り、該塩基又は核酸配列は前記増幅されるべき核酸に対
    して非−相補的であり、そして （ｃ）前記プライマーは前記増幅されるべき核酸配列の
    両端を規定し、このため１つのプライマーの伸長生成物
    がその鋳型から分離された場合に更なる合成のための鋳
    型として機能し得る、 ことを特徴とする、キット。
  2. 【請求項２】 前記特定の核酸配列がより長い配列中に
    含まれることを特徴とする、請求項１に記載の試薬キッ
    ト。
  3. 【請求項３】 前記特定の核酸配列がＤＮＡ又はＲＮ
    Ａ、例えばメッセンジャーＲＮＡであり、該ＤＮＡ又は
    ＲＮＡは１本鎖でも２本鎖でもよく、あるいは該特定の
    核酸配列はＤＮＡ−ＲＮＡハイブリッドであることを特
    徴とする、請求項１又は２に記載の試薬キット。
  4. 【請求項４】 前記特定の核酸配列がゲノムＤＮＡであ
    ることを特徴とする、請求項１〜３のいずれか１項に記
    載の試薬キット。
  5. 【請求項５】 増幅されるべき特定の核酸の各鎖につい
    てのプライマーの集合を含んで成り、該プライマーの少
    なくとも１つは前記鎖に実質的に相補的であることを特
    徴とする、請求項１〜４のいずれか１項に記載の試薬キ
    ット。
  6. 【請求項６】 プライマーが約７〜２５ヌクレオチド又
    はそれより多くのヌクレオチドを含むことを特徴とす
    る、請求項１〜５のいずれか１項に記載の試薬キット。
  7. 【請求項７】 前記非−相補的核酸配列がプロモータ
    ー、リンカー又はコード配列であることを特徴とすり、
    請求項１〜６のいずれか１項に記載の試薬キット。
  8. 【請求項８】 前記プロモーターがＴ７プロモーターで
    ある請求項７に記載の試薬キット。
  9. 【請求項９】 プライマー伸長生成物の合成を完成する
    ために機能する誘導剤をさらに含んで成ることを特徴と
    する、請求項１〜８のいずれか１項に記載の試薬キッ
    ト。
  10. 【請求項１０】 前記誘導剤がＥ．コリ（Ｅ．ｃｏｌ
    ｉ）ＤＮＡポリメラーゼＩ、Ｅ．コリＤＮＡポリメラー
    ゼＩのＫｌｅｎｏｗ断片、Ｔ４ ＤＮＡポリメラーゼ、
    他のＤＮＡポリメラーゼ、逆転写酵素、熱安定性酵素を
    含めての他の酵素から選択される酵素であることを特徴
    とする、請求項９に記載の試薬キット。
  11. 【請求項１１】 試薬キットが複数のヌクレオシドトリ
    ホスフェートをさらに含むことを特徴とする、請求項１
    〜１０のいずれか１項に記載の試薬キット。
  12. 【請求項１２】 試薬キットが容易に検出され得るヌク
    レオシドトリホスフェートをさらに含んで成ることを特
    徴とする請求項１１に記載の試薬キット。
  13. 【請求項１３】 ヘリカーゼとして知られる類からの酵
    素をさらに含んで成ることを特徴とする、請求項１〜１
    ２のいずれか１項に記載の試薬キット。
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US71697585A 1985-03-28 1985-03-28
US716975 1985-03-28
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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US4683202B1 (ja) 1990-11-27
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