JP6120718B2 - 耐病性植物 - Google Patents
耐病性植物 Download PDFInfo
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- JP6120718B2 JP6120718B2 JP2013164996A JP2013164996A JP6120718B2 JP 6120718 B2 JP6120718 B2 JP 6120718B2 JP 2013164996 A JP2013164996 A JP 2013164996A JP 2013164996 A JP2013164996 A JP 2013164996A JP 6120718 B2 JP6120718 B2 JP 6120718B2
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- 238000003757 reverse transcription PCR Methods 0.000 description 1
- 230000010153 self-pollination Effects 0.000 description 1
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Classifications
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
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- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/10—Transferases (2.)
- C12N9/12—Transferases (2.) transferring phosphorus containing groups, e.g. kinases (2.7)
- C12N9/1205—Phosphotransferases with an alcohol group as acceptor (2.7.1), e.g. protein kinases
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- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8279—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance
- C12N15/8282—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance for fungal resistance
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- C—CHEMISTRY; METALLURGY
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- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8279—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for biotic stress resistance, pathogen resistance, disease resistance
Description
HSKオルソログ配列を植物種において特定するための方法では、例えば、植物種のホモセリンキナーゼESTをデータベースにおいて特定すること;完全なホモセリンキナーゼの転写物またはcDNAを増幅するためのプライマーを設計すること;対応する完全な転写物またはcDNAを得るために、そのようなプライマーによる増幅実験を行うこと;および、転写物またはcDNAのヌクレオチド配列を決定することを含むことができる。
Van Damme他(2005、上掲)には、アブラナべと病菌に対して抵抗性である4つの変異体(dmr1−1、dmr1−2、dmr1−3およびdmr1−4)が開示される。抵抗性のレベルを、アブラナべと病菌のCala2単離体(これはE.Holub博士(Warwick HRI、Wellesbourne、UK)またはG.Van der Ackerveken博士(Department of Biology、University of Utrecht、NL)から入手可能である)を接種した7日後の実生の葉あたりの分生子柄を計数することによって調べることができる。高感受性である親系統のLer eds1−2(Parker他、1996、Plant Cell、8:2033〜2046)については、分生子柄の数が100%で設定される。感染したdmr1変異体での分生子柄形成における低下が、親系統の実生と比較して、図2に示される。
ホモセリンリン酸は、シロイヌナズナ属における、メチオニン、イソロイシンおよびトレオニンの産生における中間体である。ホモセリンキナーゼはアミノ酸の産生において重要な役割を有するので、遊離アミノ酸レベルを親系統(Ler eds1−2)および4つの異なるdmr1変異体において求めた。このために、葉全体からのアミノ酸を80%メタノールにより抽出し、その後、2回目の抽出を20%メタノールにより行った。一緒にした抽出液を乾燥し、水に溶解した。内部標準のS−アミノエチルシステイン(SAEC)を加えた後、アミノ酸をJOEL AminoTac JLC−500/V(東京、日本)におけるポストカラムニンヒドリン誘導体化による自動化されたイオン交換クロマトグラフィーによって検出した。
表1
親系統(Ler eds1−2)および変異体(dmr1−1、dmr1−2、dmr1−3およびdmr1−4)の2週齢実生の地上部分におけるホモセリン、メチオニン、トレオニンおよびイソロイシンの濃度(pmol/mg新鮮重量)。
効果がホモセリンについて特異的であるかどうかを調べるために、立体異性体のD−ホモセリンを調べた。実生全体に、水、5mMのD−ホモセリンおよび5mMのL−ホモセリンを浸透させた。天然アミノ酸のL−ホモセリンによる処理のみが、アブラナべと病菌に対する抵抗性をもたらした。水またはD−ホモセリンにより処理された実生は病原体成長における大きな低下を示さず、アブラナべと病菌に対して感受性であった。この浸透を、Cala2およびWaco9に対してそれぞれ感受性である2つのシロイヌナズナ受入体(Ler eds1−2およびWs eds1−1)に適用した。分生子柄形成をアブラナべと病菌感受性についての指標として求めた。分生子柄を、アブラナべと病菌接種後5日、および、水、D−ホモセリンまたはL−ホモセリンによる浸透後2日で計数した(図6)。L−ホモセリンによる浸透は分生子柄形成の低下およびアブラナべと病菌抵抗性を明瞭にもたらす。このことはさらに、シロイヌナズナ実生のトリパンブルー染色によって植物における病原体成長を調べることによって確認された。植物に単離体Cala2を接種した。2日後、植物を、水、5mMのD−ホモセリンおよび5mMのL−ホモセリンによる浸透によって処理した。症状を接種後5日でスコア化した。症状は、L−ホモセリン浸透の実生のみが、強く低下した病原体成長を示し、分生子柄形成を何ら示さなかったことを明瞭に示した(図7)。
1.配列相同性に基づくライブラリーのスクリーニング
シロイヌナズナのホモセリンキナーゼ遺伝子およびホモセリンキナーゼタンパク質のヌクレオチド配列およびアミノ酸配列が図8および図9に示される。
図8に示されるようなシロイヌナズナのHSK DNA配列が、標準的な分子生物学的方法を使用する任意の植物種でのDNAに対するハイブリダイゼーションによって相同的な配列について検索するためのプローブとして使用される。この方法を使用して、オルソログ遺伝子が、制限酵素により消化されたDNAに対するサザンハイブリダイゼーションによって、あるいは、ゲノムライブラリーまたはcDNAライブラリーに対するハイブリダイゼーションによって検出される。これらの技術は当業者には広く知られている。代わりのプローブとして、任意の他のより近縁な植物種のHSK DNA配列をプローブとして使用することができる。
多くの作物種については、完全なcDNA配列またはゲノム配列を続いてPCR増幅するためのプライマーを設計するために使用される部分的なHSK mRNA配列または遺伝子配列を利用することができる。5’配列および3’配列が利用可能であるときには、失われている内部配列が、HSK特異的な5’フォワードプライマーおよび3’リバースプライマーによってPCR増幅される。5’配列、内部配列または3’配列のみが利用可能であるときには、フォワードプライマーおよびリバースプライマーの両方が設計される。利用可能なプラスミドポリリンカープライマーとの組合せで、インサート物が、目的とする植物種のゲノムライブラリーおよびcDNAライブラリーから増幅される。類似した方法で、失われている5’配列または3’配列が、改良されたPCR技術(5’RACE、3’RACE、TAIL−PCR、RLM−RACEまたはベクトレットPCR)によって増幅される。
1.mRNAを、標準的な方法を使用して単離する;
2.cDNAを、オリゴdTプライマーおよび標準的な方法を使用して合成する;
3.縮重したフォワードオリゴヌクレオチドおよびリバースオリゴヌクレオチドを使用して、PCR反応を行う;
4.PCRフラグメントを標準的なアガロースゲル電気泳動によって分離し、予想されるサイズのフラグメントをゲルから単離する;
5.単離されたPCRフラグメントを、標準的な方法を使用してプラスミドベクターにクローン化する;
6.PCRによって求められるような正しいインサート物サイズを有するプラスミドをDNA配列決定によって分析する;
7.blastXを使用する配列分析により、どのフラグメントが正しい内部HSK配列を含有するかを明らかにする;
8.内部DNA配列を、その後、5’RACEおよび3’RACEのための遺伝子特異的プライマーおよび種特異的プライマーを設計して、完全なHSKコード配列を(上記で記載されたように)RLM−RACEによって得るために使用することができる。
HSKサイレンシング系統の作製がRNAiによってシロイヌナズナ属において達成されている。HSKエキソンの2つの約750bpのフラグメントを向き合う方向で含有する構築物をシロイヌナズナCol−0受入体に首尾よく形質転換した。形質転換体をアブラナべと病菌の単離体Waco9に対する抵抗性について分析した。アブラナべと病菌に対する抵抗性を与える数個の遺伝子組換え系統が得られた。HSK発現およびホモセリン蓄積の分析により、形質転換された系統において、HSK遺伝子がサイレンシングされ、アブラナべと病菌に対する抵抗性をもたらしていることが確認される。
目的とする植物種の種子を、ランダム点変異をゲノムに導入するために変異原により処理する。変異させた植物を成長させて、種子を作らせ、次世代をホモセリンの増大した蓄積についてスクリーニングする。これは、アミノ酸のホモセリンのレベルを測定することによって、HSK遺伝子の発現のレベルをモニターすることによって、または、TILLING法により、もしくは、DNA配列決定により、もしくは、ヌクレオチドの変化を特定するための任意の他の方法によりHSK遺伝子におけるミスセンス変異について探索することによって達成される。
作物への所望される変異型対立遺伝子の遺伝子移入が、交配および変異型対立遺伝子の遺伝子型決定によって達成される。これは、作物の今日でのマーカー補助育種における標準的な手法である。
表2
シロイヌナズナHSKのmRNA配列および他の植物種からのオルソログ配列の発現配列タグ(EST)およびmRNA配列についてのGI番号(GenInfo識別子)およびGenbankアクセション番号。
DMR1遺伝子座のマッピングにおいて使用された挿入/欠失(INDEL、サイズ差が括弧で示される)マーカーのプライマー配列および切断増幅多型配列(CAP、多型制限部位が括弧で示される)。
TAIRコードおよびGIコードが示される8個の候補DMR1遺伝子を増幅および配列決定するために使用されたプライマー配列
Claims (4)
- 病原体に対して抵抗性である植物であって、
前記植物および前記病原体が、レタスにおけるレタスべと病菌(Bremia lactucae)、ホウレンソウにおけるホウレンソウべと病菌(Peronospora farinosa)またはキャベツにおけるアブラナべと病菌(Hyaloperonospora parasitica)であり、
レタスべと病菌に対し抵抗性であるレタスはレタスべと病菌に対し抵抗性でないレタスと比較して増大されたホモセリンのレベルを有し、
ホウレンソウべと病菌に対し抵抗性であるホウレンソウはホウレンソウべと病菌に対し抵抗性でないホウレンソウと比較して増大されたホモセリンのレベルを有し、
アブラナべと病菌に対し抵抗性であるキャベツはアブラナべと病菌に対し抵抗性でないキャベツと比較して増大されたホモセリンのレベルを有し、
レタスべと病菌に対し抵抗性であるレタス、ホウレンソウべと病菌に対し抵抗性であるホウレンソウ、またはアブラナべと病菌に対し抵抗性であるキャベツは、コードされた酵素のホモセリンキナーゼ活性を低下させるそのホモセリンキナーゼ遺伝子に変異を有する、植物。 - ホモセリンキナーゼ遺伝子における変異が、コードされたタンパク質におけるアミノ酸置換の原因となる、請求項1に記載の植物。
- 遺伝子が、配列番号101で特定されるヌクレオチド配列および配列番号102で特定されるアミノ酸配列を有するレタス(Lactuca sativa)のホモセリンキナーゼ遺伝子である、請求項1または2に記載の植物。
- 遺伝子が、配列番号107で特定されるヌクレオチド配列および配列番号108で特定されるアミノ酸配列を有するホウレンソウ(Spinacia oleracea)のホモセリンキナーゼ遺伝子である、請求項1または2に記載の植物。
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TWI329018B (en) | 2003-05-30 | 2010-08-21 | Suntory Holdings Ltd | Anti-stress agent |
EP2175027A3 (en) | 2004-06-16 | 2010-07-07 | Rijk Zwaan Zaadteelt en Zaadhandel B.V. | Reduced susceptibility towards pathogens, in particular oomycetes, such as downy mildew in lettuce and spinach |
WO2007051483A1 (en) | 2005-11-01 | 2007-05-10 | Universiteit Utrecht Holding B.V. | Disease resistant plants |
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DK1957655T3 (en) | 2016-12-19 |
US20120278943A1 (en) | 2012-11-01 |
CN101316931A (zh) | 2008-12-03 |
ES2594886T3 (es) | 2016-12-23 |
US8354570B2 (en) | 2013-01-15 |
JP2014003980A (ja) | 2014-01-16 |
EP2522733A1 (en) | 2012-11-14 |
JP2009513152A (ja) | 2009-04-02 |
EP2514825A2 (en) | 2012-10-24 |
US8575432B2 (en) | 2013-11-05 |
JP2017093459A (ja) | 2017-06-01 |
EP1957655A2 (en) | 2008-08-20 |
PL1957655T3 (pl) | 2017-01-31 |
EP2514825A3 (en) | 2012-11-14 |
US8796511B2 (en) | 2014-08-05 |
CN101316931B (zh) | 2012-08-22 |
US20130145494A1 (en) | 2013-06-06 |
EP2522731A1 (en) | 2012-11-14 |
HK1124365A1 (en) | 2009-07-10 |
US20130333069A1 (en) | 2013-12-12 |
BRPI0618054A2 (pt) | 2011-08-16 |
US20090170703A1 (en) | 2009-07-02 |
EP2522732A1 (en) | 2012-11-14 |
WO2007051626A2 (en) | 2007-05-10 |
JP6391723B2 (ja) | 2018-09-19 |
WO2007051626A3 (en) | 2007-06-28 |
WO2007051483A1 (en) | 2007-05-10 |
EP1957655B1 (en) | 2016-08-24 |
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