JP3539961B2 - 果実特異性プロモータ - Google Patents
果実特異性プロモータ Download PDFInfo
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Description
トマトは遺伝子工学のための極めて重要な植物である。これは容易に形質転換されて外来遺伝子を発現すると共に、或る種の遺伝子により改善されることが知られた多くの特性を有する。さらに、これは多くの国で価値ある作物植物であって、米国にて販売が承認された最初のトランスジェニック食用作物である。
トマトおよび他の果実にて外来遺伝子を発現するのに有用なプロモータが知られている。たとえばトマト果実の固形物含有量は、果実特異性プロモータの背後でADPグルコース ピロホスホリラーゼ遺伝子を発現させて増加させることができる[Kishore、PCT Appl.WO 91/19806]。2A11ゲノミッククローンからのプロモータは[Pear、et al.(1989)Plant Mol.Biol.13:639−651]、トマト果実におけるADPグルコース ピロホスホリラーゼの発現を制御する。E4およびE8プロモータ[Deikman、et al(1988)The EMBO Journal 7:3315−3320]、並びにポリガラクツロナーゼに関するプロモータは果実特異性であることが知られている。しかしながら、後者の3種類はトマト果実の発育における後期段階での発現に制限され、したがって赤色果実プロモータとして知られる。2A11プロモータは初期段階の際に発現を示すが、或る種の遺伝子につき所望されるよりも弱い。したがって、緑色果実の発育に際し遺伝子を発現させる一層強力なプロモータにつきニーズが存在する。
本発明の目的はこの種のプロモータを提供することにある。さらに本発明の目的は、トマトおよび他の果実を有する作物で果実特異性として機能するプロモータを提供することにある。さらに本発明の他の目的は、これらプロモータおよび所望蛋白質または大して望ましくない蛋白質のアンチセンス配列をコードする遺伝子を有するDNA構築物を提供することにある。
発明の要点
本発明は、植物の果実体の初期発育に際し一層高いレベルにて発現を与える2種の果実特異性プロモータを提供する。これら2種のプロモータは(1)約2.3kbのDNA断片であって、その3′末端の1.4kbがSEQ ID NO:1で示されるTFM7;および(2)約900bpのDNA断片であって、その3′末端の400bpがSEQ ID NO:2で示されるTFM9である。
さらに本発明は、配列内に:
(a)TFM7およびTFM9よりなる群から選択されるプロモータと;
(b)所望蛋白質をコードするRNA配列を産生させる構造DNA配列と;
(c)植物細胞内で機能して転写終止させると共にRNA配列の3′末端にポリアデニル化ヌクレオチドを付加させる3′非翻訳領域と
を含み、前記プロモータが前記構造DNAに対し異種である組換二本鎖DNA分子をも提供する。このDNA構築物を有する植物細胞および全植物も提供される。
このDNA構築物を使用しうる植物は限定はしないがトマト、イチゴおよびマイチゴを包含する。
発明の詳細な説明
二本鎖DNA型で存在する植物遺伝子の発現はRNAポリメラーゼ酵素によるDNAの一方のストランドからのメッセンジャRNA(mRNA)の転写と、核の内側におけるmRNA一次転写物のその後の処理とを含む。この処理は、ポリアデニレート ヌクレオチドをRNAの3′末端に付加する3′非翻訳領域を含む。
DNAからmRNAへの転写は、一般にプロモータと称するDNAの領域により調節される。プロモータ領域は、RNAポリメラーゼにシグナルを与えてDNAと連携させると共に、RNAの対応する相補ストランドを作成する鋳型としてDNAストランドの一方を用いmRNAの転写を開始させる塩基配列を有する。
トマト果実の発育に際し高い特異的発現を示す新規な果実特異性プロモータが分離された。トマト果実cDNAライブラリーを用いる差次スクリーニング手法を使用して、緑色果実で特異的に発現するのに適するcDNAクローンを同定した。初期および後期発育段階にて果実から、およびトマトの葉+茎の組合せ組織から抽出されたmRNAより作成されるcDNAプローブを使用した。緑色果実にて充分に発現すると共に葉にて検出しえない発現を示すクローンを同定した。ゲノミック・サウザン分析は小(1〜2)遺伝子コピー数を示した。これらcDNAクローンのプロモータを、次いでトマトのゲノミック クローンバンクをスクリーニングすることにより分離した。これらプロモータの発現パターンを、β−グルクロニダーゼ(GUS)遺伝子への融合およびトランスジェニック果実における発育に際しGUS酵素の発現を追跡して確認した。その結果を後記実施例1に示す。
これらプロモータをWO 91/19806号に記載されたCTP−glgC16構築物に融合させた。TFM7構築物によるトマトの形質転換の結果を実施例2に示す。或いは果実における蔗糖含有量を増加させるため、ADPGPPのサブユニットの一方もしくは両方に対応するアンチセンス配列を組み込むことにより、植物ADPGPP遺伝子の作用を抑制することが望ましい。本発明によるプロモータの使用は、初期果実段階にてこれを行うのに便利な手段となりうる。
本発明のプロモータに有利に融合させうる他の遺伝子はシュークロース ホスフェートシンターゼ(SPS)を包含し、これは植物細胞におけるシュークロース生合成の全体的な速度を制御すると思われる。TFM7もしくはTFM9を動因とするSPS遺伝子の発現は、一層強度なシンク(sink)活性を有する発育果実をもたらす。
他の可能な用途はインベルターゼ遺伝子に関するものである。たとえば果実におけるようなシンク細胞でのインベルターゼの発現は、炭素利用経路(たとえば澱粉の生合成)で使用しうる代謝物まで蔗糖を分解することにより一層強力なシンクとして作用する細胞の能力を増大させるもう1つの方法である。かくして、一層多量の蔗糖がシンク組織中へ移動する。適切な組織および細胞室におけるインベルターゼの発現は、果実が強いシンクである場合、すなわち緑色果実である場合に極めて望ましい。
最後に、シュークロース シンターゼの遺伝子と共に本発明のプロモータを使用することが、SPSにつき示した理由のため望ましい。
植物形質転換/再生
本発明によるプロモータの一方を有する二本鎖のDNA分子を、任意の適する方法により植物のゲノム中に挿入することができる。適する植物形質転換ベクターはAgrobacterium tumefaciensのTiプラスミドから誘導されたもの、並びにたとえば刊行物[Herrera Estrella、et al(1983)、Nature 303:209;Klee、H.J.et al(1985)、Bio/Technology 3:637−42;並びにEPO公開第120,516号(Schilperoort et al)]により開示されたものを包含する。アグロバクテリウムのTiもしくは根誘発性(Ri)プラスミドから得られる植物形質転換ベクターの他に、別の方法を用いて本発明のDNA構築物を植物細胞中に挿入することもできる。この種の方法はたとえばリポソーム、電子穿孔法、フリーDNA取込みを増加させる薬品、ミクロプロジェクチル・ボンバードメントによるフリーDNA供給およびウィルスもしくは花粉を用いる形質転換を包含する。
双子葉植物の形質転換に使用するための特に有用なアグロバクテリウム系植物形質転換ベクターはプラスミドベクターpMON505[Rogers,S.G.et al.(1987)“Improved Vectors for Plant Transformation:Expression Cassette Vectors and New Selectable Markers"in Methods in Enzymology,ed.Wu and Grossman,pp 253−277,San Diego:Academic Press)]である。二成分ベクターpMON505は、Tiプラスミドのホモロジー領域(LIH)がminiRK2プラスミドのSma I断片、すなわちpTJS75に対し3.8kb Hind IIIで置換されたpMON200の誘導体である[Schmidhauser、T.J.およびD.R.Helinski(1985)、J.Bacteriol.164−155]。この断片は複数のPK2オリジン(すなわちoriV)を有し、さらに三親交配過程を用いアグロバクテリウムに結合させるための移動のオリジン(すなわちoriT)をも有する[Horsch、R.B.およびH.Klee(1986)、PNAS U.S.A.83:4428−32]。プラスミドpMON505は、所望DNA断片を挿入するための合成マルチリンカー、植物細胞におけるカナマイシン耐性のためのキメラNOS/NPTII/NOS遺伝子、大腸菌およびA.ツメファシエンスにて選択するためのスペクチノマイシン/ストレプトマイシン耐性決定基、形質転換体を容易に評価するための完全ノパリンシンターゼ遺伝子および後代における遺伝形質、並びに大腸菌における多量のベクターの作成を容易化させる複製のpBR322オリジンを含めpMON200の重要な特徴を全て保持する。プラスミドpMON505は、pTiT37ノパリン型T−DNAの右側末端から誘導された単一のT−DNAボーダーを有する。サウザン分析は、プラスミドpMON505およびこれらを有するDNAが植物ゲノムに組み込まれること、すなわち全プラスミドは植物ゲノム中に挿入されたT−DNAであることを示す。組み込まれたDNAの一方の末端は右側ボーダー配列とノパリンシンターゼ遺伝子との間に位置し、他方の末端はボーダー配列とpBR322配列との間に位置する。
他の特に有用なTiプラスミド カセットベクターはpMON17227である。このベクターはバリー等によりWO 92/04449号に記載され、多くの植物につき優秀な選択マーカー遺伝子であるグリホセート耐性を付与する酵素をコードする遺伝子を含む。
本発明のプロモータにより被動される選択遺伝子を含む充分な個数の細胞(もしくは原形質)が得られれば、これら細胞(または原形質)を全植物に再生させる。再生工程に関する方法の選択は重要でなく、適する手法をトマトおよびコショウからの宿主につき用いることができる。
以下、本発明の範囲を限定することなく実施例により本発明の実施につき一層詳細に説明する。すなわち本発明の思想および範囲を逸脱することなく本発明の方法および遺伝子につき種々の改変をなしうることが当業者には了解されよう。
実施例1
上記したTFM7およびTFM9と称する2種の緑色果実プロモータを、Lycopersicon esculentum cv.VF36ゲノムライブラリーから分離すると共に特徴付けした。これらのそれぞれにつき、プロモータを誘導した5′末端、非翻訳およびプロモータ領域の部分配列をここに示す。SEQ ID NO:1はTFM7のものである。この2.3kbプロモータ断片は5′末端として内部XBal部位を有し、推定翻訳開始点まで延びる(後者の点にBgl II認識部位を入れて改変される)。SEQ ID NO:2はTFM9のものである。〜900bp TFM9プロモータ断片は内部Sal I部位から推定翻訳開始点まで延びる(後者の点にBgl II認識部位を入れて改変)。
これらプロモータのそれぞれをGUS遺伝子に融合させ、トマトに形質転換させた。再生されたトマト植物をそのライフサイクル全体にわたりGUSの発現を証明するため観察した。その結果を第1表および第2表に示す。そこに示した全ての数値は、トランスジェニック系列からの3種もしくは4種のR1 GUS陽性植物から収穫された少なくとも3個の果実からの標準誤差を含む平均値を示す。数値のフロリダーダ野生型比較値物からのGUS活性読値をも示す。発育段階および組織段階はインマチュア・グリーン1(2〜3cm果実)、インマチュア・グリーン2(4〜5cm果実)、マチュア・グリーン果実(少なくとも2個の子房室が満たされる)、ターニング(果実の10〜20%が桃色もしくは赤色である)、および若葉である。「ND」は、検出しうる単位が記録されなかったことを意味する。第1表において、トランスジェニック系列11541、11420および11305はpTFM7/GUS/nos構築物を有する選択系列である。第2表において、トランスジェニック系列11256、11269および11290はpTFM7/GUS/nos構築物を有する選択系列である。
実施例2
TFM7およびTFM9をWO 91/19806号に開示されたキメラCTP−GlgC16遺伝子(および適する3′配列)に融合させ、発現カセットを下記するように植物形質転換ベクター中に移動させた。
TFM7プロモータを操作を容易化すべくベクターpMON999に結合させて、プラスミドpMON16987を生ぜしめた。CTP−glgC16キメラ遺伝子との融合は、pMON16987からのHind III−Bgl II TFM7プロモータ断片とpMON20102からのBgl II−Sac I断片(WO 91/19806号に開示されたキメラ遺伝子を有する)とのトリプル結合により得られ、これをHind IIIおよびSac Iで消化された2成分植物形質転換ベクターpMON10098(WO 91/19806号の第11図参照)に入れて行った。このプラスミドpMON16989を次いで、トマト品種UC204Cを形質転換させるべく用いた。
TFM9プロモータを実質的に上記したようにCTP1−glgC16に融合させた。Sal I−BamH I TFM9プロモータ断片+GUSを同じ酵素で切断したpEMBL18+に結合させ、pMON22701を得た。次いでTFM9プロモータをHind III−Bgl II断片として除去し(pMON22701から)、pMON20102からのCTP1−glgC16 Bgl II−Sac I断片とHind III−SacI断片pMON10098に結合させて、pMON22709を得た。このプラスミドを用いてトマト品種UC204Cを形質転換させた。
トマト植物細胞を刊行物[McCormick et al.(1986)]に記載された方法によりアグロバクテリウム菌株を用いて形質転換させた。特に、7〜8日令の苗から子葉を得た。種子を次の手順により表面殺菌した:(1)種子を水中に15分間にわたり浸漬する;(2)70%EtOH中に1分間にわたり浸漬し、次いで無菌水で洗浄する;(3)1NのNaOHに20分間浸漬する;(4)無菌水で2回洗浄する;(5)ツイーン20を含む25%クロロックスに25分間間浸漬する;(6)無菌脱イオン水で3回洗浄する。これら種子をフィタ・トレー(シグマ社)にて25mg/Lのアスコルビン酸を添加した上記のデイビス発芽培地で発芽させた。種子を2〜3日間にわたり暗所内で28℃にて培養し、次いで成長室内で25℃および40%湿度にて80アインシュタインm-2S-1の強度を有する冷白色光の下に成長させた。光周期は16時間の明所および8時間の暗所とした。
発芽を開始してから7〜8日間の後、子葉を上記のように移植した。子葉をカルジーン培地+アセトシリンゴンおよび1mMのガラクツロン酸(抗体を含有せず)で構成された「フィーダー・プレート」にて上記した条件下で予備培養した。
次いで子葉に上記プラスミドを有するアグロバクテリウムの対数期溶液を接種した。アグロバクテリウムの濃度は約5×108細胞/mLである。子葉を細菌溶液に8分間にわたり浸漬させ、次いで無菌ワットマン フィルタ円盤にブロットして過剰の溶液を除去し、次いで最初のフィーダープレートに再び載せて2〜3日間にわたり一緒に培養した。
子葉を2mg/Lのゼアチンリボシドと500μg/mLのカルベニシリンと100μg/mLのカナマイシンとを含むデイビス再生培地を含有した選択プレートに移した。2〜3週間の後、カルスおよび/またはシュートの形成を有する子葉を、同レベルでカルベニシリンおよびカナマイシンを含有する新たなデイビス再生プレートに移した。実験をこの時点で形質転換体につき評価した。カルス組織を定期的な3週間間隔でサブ培養すると共に、異常な構造体をトリミングして発育シュート芽が再生し続けるようにした。シュートは3〜4か月以内で発育した。
シュートが発育した後、これらをカルス組織から綺麗に切除し、発根選択プレートに移植した。これらプレートは、50μg/mLのカナマイシンと500μg/mLのカルベニシリンとを含有する0.5XMSOを含む。これらシュートは2週間以内に選択培地にて根を形成する。2週間後にシュートが出現しなければ、シュートをトリミングして選択培地に再移植する。シュート培養物を22℃の温度にてパーシバル(percivals)にて培養する。次いで根を伴うシュートを根が長さ約2cmになった際に鉢植えした。これら植物を18時間の光および6時間の暗所の光周期にて2〜3週間にわたり21℃の成長室内で栽培した後、温室に移した。温室内で、植物を日中は26℃かつ夜間は21℃の温度にて成長させた。光周期は13時間の光および11時間の暗所として成熟させた。
pMON16989(TFM7プロモータ)で形質転換された植物からの果実が得られ、これを試験した。TFM7はglgC16酵素の高発現を緑色果実にて生ぜしめた(>0.1%抽出蛋白質)が、glgC16発現は極めて弱いか或いは熟成果実では検出しえない。TFM7は或る系列における成熟果実にて澱粉を生成するのに対し、比較は常に“0"の沃素尺度を示し、これは澱粉が存在しないことを示す。これら果実からのジュースは極めて粘性であり、可溶性固形物が多くの系列にて増加する。比較データを第3表に示す。可溶性固形物および澱粉の量を熱処理トマトジュースからのセルムにて測定した。1滴の沃素溶液を濾過セルムに添加すると共に色強度を0〜4の尺度(この尺度にて黄色=0(澱粉なし)〜暗青色=4(高含有量の澱粉)である)にて測定することにより、澱粉を測定した。
配列リスト
(1)一般的情報:
(i)出願人:
(A)名称:モンサント・カンパニー
(B)町名:800ノース・リンドバーグ・ボウレバード
(C)市名:セントルイス
(D)州名:ミズーリ
(E)国名:アメリカ合衆国
(F)郵便コード(ZIP):63167
(G)電話番号:(314)694−3131
(H)テレファックス:(314)694−5435
(ii)発明の名称:果実特異性プロモータ
(iii)配列の数:2
(iv)コンピュータ解読フォーム:
(A)メディア種類:フロッピー・ディスク
(B)コンピュータ:IBM PCコンパチブル
(C)作動システム:PC−DOS/MS−DOS
(D)ソフトウエア:パテント・レリースNo.1.0、バージョンNo.1.25(EPO)
(vi)先行出願データ:
(A)出願番号:US 08/090523
(B)出願日:12−JUL−1993
(2)SEQ ID NO:1の情報:
(i)配列特性:
(A)長さ:1478塩基対
(B)種類:核酸
(C)鎖型:二本鎖
(D)トポロジー:線状
(ii)分子型:DNA(ゲノミック)
(xi)配列説明:SEQ ID NO:1:
(2)SEQ ID NO:2の情報:
(i)配列特性:
(A)長さ:450塩基対
(B)種類:核酸
(C)鎖型:二本鎖
(D)トポロジー:線状
(ii)分子型:DNA(ゲノミック)
(xi)配列説明:SEQ ID NO:2:
Claims (11)
- 3'末端が配列番号1で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる2.3kbの単離された果実特異性プロモータ。
- 3'末端が配列番号2で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる900bpの単離された果実特異性プロモータ。
- 配列内に:
(a)3'末端が配列番号1で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる2.3kbDNAフラグメントのTFM7、及び3'末端が配列番号2で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる900bpDNAフラグメントのTFM9からなる群から選択されるプロモータと;
(b)所望蛋白質をコードするRNA配列を産生させる構造DNA配列と;
(c)植物細胞内で機能して転写終止させると共にRNA配列の3′末端にポリアデニル化ヌクレオチドを付加させる3′非翻訳領域と
を含み、前記プロモータが前記構造DNAに対し異種であることを特徴とする組換二本鎖DNA分子。 - DNA配列がADPグルコース ピロホスホリラーゼをコードする請求項3に記載のDNA分子。
- 前記酵素がglgC16である請求項4に記載のDNA分子。
- 配列内に:
(a)3'末端が配列番号1で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる2.3kbDNAフラグメントのTFM7、及び3'末端が配列番号2で表されるDNA配列であるリコペルシコン エスカレンタム(Lycopersicon esculentum)から得られる900bpDNAフラグメントのTFM9よりなる群から選択されるプロモータと;
(b)所望蛋白質をコードするRNA配列を産生させる構造DNA配列と;
(c)植物細胞内で機能して転写終止させると共にRNA配列の3′末端にポリアデニル化ヌクレオチドを付加させる3′非翻訳領域と
を含み、前記プロモータが前記構造DNAに対し異種であることを特徴とする組換二本鎖DNA分子を含む植物細胞。 - DNA配列がADPグルコース ピロホスホリラーゼをコードする請求項6に記載の植物細胞。
- 前記酵素がglgC16である請求項7に記載の植物細胞。
- 細胞がトマトからの細胞である請求項6に記載の植物細胞。
- 請求項9に記載の植物細胞よりなるトマト植物。
- DNA配列がADPグルコース ピロホスホリラーゼをコードする請求項10に記載のトマト植物。
Applications Claiming Priority (3)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
US090,523 | 1993-07-12 | ||
US08/090,523 US5498830A (en) | 1990-06-18 | 1993-07-12 | Decreased oil content in plant seeds |
PCT/US1994/007072 WO1995002696A1 (en) | 1993-07-12 | 1994-06-27 | Fruit specific promoters |
Publications (2)
Publication Number | Publication Date |
---|---|
JPH09500274A JPH09500274A (ja) | 1997-01-14 |
JP3539961B2 true JP3539961B2 (ja) | 2004-07-07 |
Family
ID=22223158
Family Applications (1)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
JP50456895A Expired - Fee Related JP3539961B2 (ja) | 1993-07-12 | 1994-06-27 | 果実特異性プロモータ |
Country Status (12)
Country | Link |
---|---|
US (4) | US5498830A (ja) |
EP (2) | EP0708835B1 (ja) |
JP (1) | JP3539961B2 (ja) |
AT (2) | ATE223969T1 (ja) |
AU (1) | AU675158B2 (ja) |
CA (1) | CA2165669C (ja) |
DE (2) | DE69431352T2 (ja) |
DK (1) | DK0634491T3 (ja) |
ES (1) | ES2148308T3 (ja) |
GR (1) | GR3034038T3 (ja) |
PT (1) | PT634491E (ja) |
WO (1) | WO1995002696A1 (ja) |
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GR3034038T3 (en) | 2000-11-30 |
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ATE223969T1 (de) | 2002-09-15 |
US5608149A (en) | 1997-03-04 |
EP0634491A1 (en) | 1995-01-18 |
EP0634491B1 (en) | 2000-05-24 |
ATE193328T1 (de) | 2000-06-15 |
WO1995002696A1 (en) | 1995-01-26 |
DK0634491T3 (da) | 2000-09-04 |
DE69431352T2 (de) | 2003-05-08 |
AU7316094A (en) | 1995-02-13 |
US5608150A (en) | 1997-03-04 |
AU675158B2 (en) | 1997-01-23 |
US6538179B1 (en) | 2003-03-25 |
ES2148308T3 (es) | 2000-10-16 |
CA2165669A1 (en) | 1995-01-26 |
EP0708835A1 (en) | 1996-05-01 |
PT634491E (pt) | 2000-11-30 |
US5498830A (en) | 1996-03-12 |
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