JP5818114B2 - 果実特異的プロモーター - Google Patents
果実特異的プロモーター Download PDFInfo
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- JP5818114B2 JP5818114B2 JP2013506945A JP2013506945A JP5818114B2 JP 5818114 B2 JP5818114 B2 JP 5818114B2 JP 2013506945 A JP2013506945 A JP 2013506945A JP 2013506945 A JP2013506945 A JP 2013506945A JP 5818114 B2 JP5818114 B2 JP 5818114B2
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Description
トマト(Solanum lycopersicum)品種マイクロトム(Micro-Tom)のESTマイクロアレイに対してマイクロトムの緑熟果実から抽出したmRNA由来のプローブをハイブリダイズさせて得られたマイクロアレイデータに基づき、高発現が認められた遺伝子を候補緑熟果実高発現遺伝子として選抜した。選抜された候補遺伝子を、ID番号LA15CA04、LA22CD07、LC09AH08、LC04DC11、LA12AA05、LA14AD08及びFB14DB02で示す。
実施例1で選抜した候補遺伝子について、RT-PCR発現解析を行った。まず、生育3ヶ月目のトマト(Solanum lycopersicum)の品種マイクロトム(Micro-Tom)の生物試料から、RT-PCRに使用するトータルRNAを、以下の方法により抽出した。まずマイクロトムの葉、花、茎、根、緑熟果実、及び赤熟果実を約1 gずつ採取し、それぞれ独立に液体窒素で凍結させた。凍結させた各組織を独立に乳鉢中で破砕し、粉状にした。それぞれの粉状サンプルを50 mlプラスチックチューブに移し、これらのチューブに約10 mlのTRIzol(R)(Invitrogen, USA)を加え、ボルテックスで2〜3分間混合した。これらを室温で10分間インキュベート後、各チューブに1 mlのクロロフォルムを加えボルテックスで2〜3分間混合した。室温で1分間インキュベート後、10000 rpmで10分間、4℃で遠心分離を行った。各チューブの上層(水層)をそれぞれ新しい50mlプラスチックチューブに移し、等量のフェノール/クロロフォルム/イソアミルアルコール溶液を加え、ボルテックスにより2〜3分間混合した。室温で各チューブを2〜3分間インキュベート後、10000 rpmで10分間、4℃で遠心分離を行った。各チューブの上層(水層)をそれぞれ新しい50 mlプラスチックチューブに移し、等量のフェノール/クロロフォルム溶液を加え、ボルテックスにより2〜3分間混合した。室温で各チューブを2〜3分間インキュベート後、10000 rpmで10分間遠心分離を行った。各チューブの上層(水層)をそれぞれ新しい50 mlプラスチックチューブに移し、これらに等量のイソプロパノールを加え、ボルテックスで2〜3分間混合した。室温で5分間インキュベート後、10000 rpmで10分間、4℃で遠心分離を行った。上清を除去し、ペレットに-20℃で冷却した70%エタノールを加え、ペレットを洗浄した。10000 rpmで10分間、4℃で遠心分離を行った後、上清を除去した。ペレットを室温で5〜10分間インキュベートすることによりエタノールを完全に除去した後、RNase及びDNaseを含まない滅菌水でペレットを溶解し、RNA溶液とした。このRNA溶液について、マイクロプレートリーダーSafire(Tecan, Switzerland)を使用し、OD260値を測定しRNA量を算出した。次いでDNase処理した1μgのトータルRNAを鋳型として、ポリTプライマーとSuperScript(R) II(Invitrogen, USA)を使用して、逆転写反応によりファーストストランドcDNAを作製した。
上記候補遺伝子の機能を調べる目的で、NCBI(National Center for Biotechnology Information, U.S.A.)のBLASTNプログラムを用いて配列解析を行った。
プロモーター解析が行われていない5つの遺伝子LA22CD07、LA12AA05、LA14AD08、Les.3122.2.A1_a_at、及びLesAffx.6852.1.S1_atのそれぞれから、以下のようにしてプロモーター領域の単離を試みた。
ゲノムDNAは、トマト(品種マイクロトム又はマネーメーカー)の葉から、臭化セチルトリメチルアンモニウム(ナカライテスク)を使用したCTAB法(Murray and Thompson, 1980)によって抽出した。具体的には、まず約3 gの葉を液体窒素下で乳鉢中で破砕し、粉状にした。この粉末を、70℃の5 mlの2xCTAB液の入ったプラスチックチューブへ移し、60分間、55℃でゆっくり振盪した。このチューブに、5 mlのクロロホルム/イソアミルアルコールを加え、30分間、室温でゆっくり振盪した。次に、5000 rpmで15分間、室温で遠心した後、上層(水層)を新しいプラスチックチューブに移し、1/10容量の10% CTAB液を加え、混合した。10分間、室温でゆっくり振盪後、5000 rpmで15分間、室温で遠心した。上層(水層)を新しいチューブに移し、等量の沈殿用バッファーを加え、転倒混和し30分間、室温でインキュベートした。このチューブを5000 rpmで15分間、室温で遠心した後、上清を除去した。ペレットに5 mlの1M NaCl-TE溶液(RNaseIを含む)を加え、55℃でペレットを溶解させた。これに5 mlのイソプロパノールを加え、転倒混和し、室温で30分間インキュベートした。これを、5000 rpmで15分間、室温で遠心した後、上清を除去した。ペレットに5 mlの70%エタノールを加え、転倒混和した後、5000 rpmで15分間、室温で遠心した。上清を除去した後、ペレットを室温で軽く乾燥し、TE溶液に溶解した。マイクロプレートリーダーSafire(Tecan, Switzerland)を使用し、OD260値を測定し、ゲノムDNA量を算出した。
ゲノムDNAを鋳型として用いて、Genome WalkerTM Universal Kit(Clonetec, USA)を使用したゲノムウォーキング法により、各遺伝子の上流域を増幅した。具体的には、最初に、上記で調製したマイクロトム由来のゲノムDNAを制限酵素Dra I、EcoR V、 Pvu II又はStu Iで消化し、得られた切断断片の両端にライゲーションによりアダプター配列を連結して、4種のゲノムライブラリーを作製した。次に、各ゲノムライブラリーを鋳型にして、アダプタープライマー1(AP1)と遺伝子特異的プライマー1(GSP1)で一次PCRを行った。PCR反応は総量50 μlで行った。その反応液組成は、各プライマー20 pM、ゲノムライブラリー1 μl、10x PCRバッファー 5 μl、dNTPs 4 μl、ポリメラーゼ酵素1.0 μl、及び滅菌水37.0 μlであった(10x PCRバッファー、dNTPs、ポリメラーゼ酵素はExpand High Fidelity PCR システム(Roche)由来のものを使用)。PCR反応は、94℃で0.25分の変性、72℃で3分のアニーリング及び伸長を1サイクルとして7サイクル行った後、94℃で0.25分の変性、67℃で3分のアニーリング及び伸長を32サイクル行い、最後に67℃で7分の伸長を行う熱サイクル条件で実施した。次に、一次PCRの反応産物を鋳型として、アダプタープライマー2(AP2)と遺伝子特異的プライマー2(GSP2)を使用して二次PCRを行った。反応液組成及び反応条件は、一次PCRと同様である。得られたPCR産物 5 μlをアガロースゲルで電気泳動し、増幅産物を確認した。
(1)植物発現ベクターの構築及びアグロバクテリウムへの遺伝子導入
プロモーター活性の評価は、GUS遺伝子発現活性に基づいて行った。まず、GUS遺伝子を含む植物発現用ベクターであるpBI121に、各遺伝子のプロモーター領域を含むDNA断片を以下のようにして導入した。実施例4でpCR(R)-Blunt II-TOPO(R)ベクターに組み込んだプロモーター領域を制限酵素処理により切り出し、切り出したDNA断片を、35Sプロモーターを除去した発現ベクターpBI121のGUS遺伝子上流部位にライゲーション反応により組み込んだ。作製したベクターをエレクトロポレーションによりAgrobacterium tumefaciens GV3101株に導入し、組換えアグロバクテリウムを、抗生物質カナマイシンを50 mg/Lで添加したLB寒天培地上で選抜した。
トマトの緑熟果実を使用した一過性発現解析を、Orzaezら(2006)の方法に従って行った。具体的には、上記で作製した組換えアグロバクテリウムを、抗生物質カナマイシンを50 mg/Lで添加した5 mlの液体YEB培地で28℃で一晩培養した後、抗生物質カナマイシンを50 mg/Lで添加した50 mlの誘導培地(0.5% 肉エキス(Beef extract)、0.1% 酵母エキス、0.5% ペプトン、0.5% スクロース、2 mM MgSO4, 20 μM アセトシリンゴン、10 mM MES、pH 5.6)でさらに一晩培養した。この培養液を3000 rpmで10分間室温で遠心し、集菌した。集菌したペレットを感染培地(10 mM MgCl2、10 mM MES、200 μM アセトシリンゴン、pH5.6)にOD600=1.0となる濃度で懸濁し、2時間、室温でゆっくり浸透させた。この菌液500 μlを1 mlシリンジに移し、菌液を含むシリンジの針を、マイクロトムから切り取った緑熟果実に刺し、菌液を果実に注入した。この果実を、2 mlの蒸留水に浸した濾紙を含む9センチプラスチックシャーレに入れ、25℃、16時間日長で4日間インキュベートした。
実施例5(1)で、果実での活性が確認された遺伝子(LA22CD07、Les.3122.2.A1_a_at、及びLesAffx.6852.1.S1_at)の各プロモーター領域を含むベクターをA. tumefaciens GV3101株に導入して得た組換えアグロバクテリウムを用いて、Sunら(Plant Physiol., 114:1547-1556, 2006)の方法により、マイクロトムの形質転換体を作製した。また対照として、35Sプロモーターの制御下にGUS遺伝子を含む発現ベクターpBI121を導入した組換えアグロバクテリウムを用いて、同様にマイクロトムの形質転換体を作製した。
配列番号2:LesAffx.6852.1.S1_atプロモーター
配列番号3〜36:プライマー
Claims (8)
- 配列番号1又は2で示される塩基配列に対して90%以上の同一性を有する塩基配列からなり、かつ緑熟果実においてプロモーター活性を有する果実特異的プロモーターDNA。
- 請求項1に記載の果実特異的プロモーターDNAを含む、発現ベクター。
- 前記果実特異的プロモーターDNAの下流に連結された遺伝子をさらに含む、請求項2に記載の発現ベクター。
- 請求項1に記載の果実特異的プロモーターDNAとその下流に連結された遺伝子とを含む、DNA構築物。
- 請求項2若しくは3に記載の発現ベクター又は請求項4に記載のDNA構築物を含む、形質転換細胞。
- 請求項3に記載の発現ベクター又は請求項4に記載のDNA構築物を導入した、形質転換植物。
- 請求項3に記載の発現ベクター又は請求項4に記載のDNA構築物を植物細胞に導入し、形質転換植物を育成して果実を形成させ、果実中の前記遺伝子の発現を確認することを含む、形質転換植物の作製方法。
- 請求項6に記載の形質転換植物を育成して果実を形成させ、発現された遺伝子産物を果実から取得することを含む、遺伝子産物の組換え生産方法。
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JPN6011019817; DEIKMAN J., et al., "Interaction of a DNA binding factor with the 5'-flanking region of an ethylene- * |
JPN6011019817; DEIKMAN J., et al., Interaction of a DNA binding factor with the 5'-flanking region of an ethylene-r * |
JPN6015016376; YANO, Kentaro et al.: '"MiBASE: A database of a miniature tomato cultivar Micro-Tom"' Plant Biotechnology Vol. 23, 2006, pp. 195-198 * |
JPN6015016378; OZAKI, Soichi et al.: '"Coexpression Analysis of Tomato Genes and Experimental Verification of Coordinated Expression of Ge' DNA Research Vol. 17, 2010, pp. 105-116 * |
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