JP6891198B2 - プロトポルフィリノーゲンオキシダーゼまたはその変異体を用いる除草剤耐性付与および/または増進のための組成物および方法 - Google Patents
プロトポルフィリノーゲンオキシダーゼまたはその変異体を用いる除草剤耐性付与および/または増進のための組成物および方法 Download PDFInfo
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- CONWAEURSVPLRM-UHFFFAOYSA-N lactofen Chemical compound C1=C([N+]([O-])=O)C(C(=O)OC(C)C(=O)OCC)=CC(OC=2C(=CC(=CC=2)C(F)(F)F)Cl)=C1 CONWAEURSVPLRM-UHFFFAOYSA-N 0.000 description 1
- 150000002632 lipids Chemical class 0.000 description 1
- 238000011068 loading method Methods 0.000 description 1
- PWPJGUXAGUPAHP-UHFFFAOYSA-N lufenuron Chemical compound C1=C(Cl)C(OC(F)(F)C(C(F)(F)F)F)=CC(Cl)=C1NC(=O)NC(=O)C1=C(F)C=CC=C1F PWPJGUXAGUPAHP-UHFFFAOYSA-N 0.000 description 1
- 239000011777 magnesium Substances 0.000 description 1
- 229910052749 magnesium Inorganic materials 0.000 description 1
- 108010086470 magnesium chelatase Proteins 0.000 description 1
- 210000001161 mammalian embryo Anatomy 0.000 description 1
- 230000013011 mating Effects 0.000 description 1
- 238000005259 measurement Methods 0.000 description 1
- MJGFBOZCAJSGQW-UHFFFAOYSA-N mercury sodium Chemical compound [Na].[Hg] MJGFBOZCAJSGQW-UHFFFAOYSA-N 0.000 description 1
- 230000004060 metabolic process Effects 0.000 description 1
- 229930182817 methionine Natural products 0.000 description 1
- 230000011987 methylation Effects 0.000 description 1
- 238000007069 methylation reaction Methods 0.000 description 1
- 238000013508 migration Methods 0.000 description 1
- 230000005012 migration Effects 0.000 description 1
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- 235000010446 mineral oil Nutrition 0.000 description 1
- 210000003470 mitochondria Anatomy 0.000 description 1
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- 238000001823 molecular biology technique Methods 0.000 description 1
- 239000002808 molecular sieve Substances 0.000 description 1
- 231100000350 mutagenesis Toxicity 0.000 description 1
- 229960004927 neomycin Drugs 0.000 description 1
- 238000006386 neutralization reaction Methods 0.000 description 1
- 229910052759 nickel Inorganic materials 0.000 description 1
- 238000010899 nucleation Methods 0.000 description 1
- 239000002773 nucleotide Substances 0.000 description 1
- 125000003729 nucleotide group Chemical group 0.000 description 1
- 235000016709 nutrition Nutrition 0.000 description 1
- 235000014571 nuts Nutrition 0.000 description 1
- 239000001301 oxygen Substances 0.000 description 1
- 229910052760 oxygen Inorganic materials 0.000 description 1
- 238000010422 painting Methods 0.000 description 1
- 239000002540 palm oil Substances 0.000 description 1
- 239000002245 particle Substances 0.000 description 1
- 235000020232 peanut Nutrition 0.000 description 1
- 239000000137 peptide hydrolase inhibitor Substances 0.000 description 1
- 235000011197 perejil Nutrition 0.000 description 1
- 239000013500 performance material Substances 0.000 description 1
- YQJNPNXZUFSGOS-UHFFFAOYSA-N phenyl 4,4-difluoropiperidine-1-carboxylate Chemical compound FC1(CCN(CC1)C(=O)OC1=CC=CC=C1)F YQJNPNXZUFSGOS-UHFFFAOYSA-N 0.000 description 1
- 125000001997 phenyl group Chemical group [H]C1=C([H])C([H])=C(*)C([H])=C1[H] 0.000 description 1
- 239000010452 phosphate Substances 0.000 description 1
- 230000000243 photosynthetic effect Effects 0.000 description 1
- 125000000587 piperidin-1-yl group Chemical group [H]C1([H])N(*)C([H])([H])C([H])([H])C([H])([H])C1([H])[H] 0.000 description 1
- DVMSBIVGIAGNNI-UHFFFAOYSA-N piperidin-1-ylcarbamic acid Chemical compound OC(=O)NN1CCCCC1 DVMSBIVGIAGNNI-UHFFFAOYSA-N 0.000 description 1
- 230000008635 plant growth Effects 0.000 description 1
- 230000037039 plant physiology Effects 0.000 description 1
- 229920003023 plastic Polymers 0.000 description 1
- 229920001223 polyethylene glycol Polymers 0.000 description 1
- 239000000256 polyoxyethylene sorbitan monolaurate Substances 0.000 description 1
- 235000010486 polyoxyethylene sorbitan monolaurate Nutrition 0.000 description 1
- 239000002244 precipitate Substances 0.000 description 1
- 238000001556 precipitation Methods 0.000 description 1
- 238000002360 preparation method Methods 0.000 description 1
- 210000001236 prokaryotic cell Anatomy 0.000 description 1
- TURAMGVWNUTQKH-UHFFFAOYSA-N propa-1,2-dien-1-one Chemical group C=C=C=O TURAMGVWNUTQKH-UHFFFAOYSA-N 0.000 description 1
- 230000004952 protein activity Effects 0.000 description 1
- 230000012846 protein folding Effects 0.000 description 1
- 239000012264 purified product Substances 0.000 description 1
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- 230000002829 reductive effect Effects 0.000 description 1
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- 239000011347 resin Substances 0.000 description 1
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- 238000005204 segregation Methods 0.000 description 1
- 230000035945 sensitivity Effects 0.000 description 1
- 238000002864 sequence alignment Methods 0.000 description 1
- URGAHOPLAPQHLN-UHFFFAOYSA-N sodium aluminosilicate Chemical compound [Na+].[Al+3].[O-][Si]([O-])=O.[O-][Si]([O-])=O URGAHOPLAPQHLN-UHFFFAOYSA-N 0.000 description 1
- 229910001023 sodium amalgam Inorganic materials 0.000 description 1
- 238000002415 sodium dodecyl sulfate polyacrylamide gel electrophoresis Methods 0.000 description 1
- 239000007787 solid Substances 0.000 description 1
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- 230000001225 therapeutic effect Effects 0.000 description 1
- 229960004659 ticarcillin Drugs 0.000 description 1
- OHKOGUYZJXTSFX-KZFFXBSXSA-N ticarcillin Chemical compound C=1([C@@H](C(O)=O)C(=O)N[C@H]2[C@H]3SC([C@@H](N3C2=O)C(O)=O)(C)C)C=CSC=1 OHKOGUYZJXTSFX-KZFFXBSXSA-N 0.000 description 1
- UZKQTCBAMSWPJD-UQCOIBPSSA-N trans-Zeatin Natural products OCC(/C)=C\CNC1=NC=NC2=C1N=CN2 UZKQTCBAMSWPJD-UQCOIBPSSA-N 0.000 description 1
- UZKQTCBAMSWPJD-FARCUNLSSA-N trans-zeatin Chemical compound OCC(/C)=C/CNC1=NC=NC2=C1N=CN2 UZKQTCBAMSWPJD-FARCUNLSSA-N 0.000 description 1
- 230000002103 transcriptional effect Effects 0.000 description 1
- 238000010361 transduction Methods 0.000 description 1
- 230000026683 transduction Effects 0.000 description 1
- OUYCCCASQSFEME-UHFFFAOYSA-N tyrosine Natural products OC(=O)C(N)CC1=CC=C(O)C=C1 OUYCCCASQSFEME-UHFFFAOYSA-N 0.000 description 1
- 238000000108 ultra-filtration Methods 0.000 description 1
- 235000013311 vegetables Nutrition 0.000 description 1
- 229910052902 vermiculite Inorganic materials 0.000 description 1
- 239000010455 vermiculite Substances 0.000 description 1
- 235000019354 vermiculite Nutrition 0.000 description 1
- 239000011534 wash buffer Substances 0.000 description 1
- 229940023877 zeatin Drugs 0.000 description 1
- DGVVWUTYPXICAM-UHFFFAOYSA-N β‐Mercaptoethanol Chemical compound OCCS DGVVWUTYPXICAM-UHFFFAOYSA-N 0.000 description 1
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Description
配列番号2のポリペプチド、配列番号2のポリペプチドの変異体、配列番号4のポリペプチド、配列番号4のポリペプチドの変異体、およびこれらと95%以上、98%以上、または99%以上の配列相同性を有するポリペプチド;
前記ポリペプチド、ポリペプチド変異体、およびこれらと95%以上、98%以上、または99%以上の配列相同性を有するポリペプチドを暗号化するポリヌクレオチド;
前記ポリヌクレオチドを含む組換えベクター;および
前記組換えベクターを含む組換え細胞
からなる群より選択された1種以上を含む、植物および/または藻類の除草剤に対する耐性付与および/または増進用組成物を提供する。一例で、前記配列番号2のポリペプチドを暗号化するポリヌクレオチドは配列番号1のポリヌクレオチド配列を含むものであってもよく、配列番号4のポリペプチドを暗号化するポリヌクレオチドは配列番号3のポリヌクレオチド配列を含むものであってもよいが、これに制限されるのではない。
配列番号2のポリペプチド、配列番号2と95%以上、98%以上、または99%以上の配列相同性を有するポリペプチド、配列番号2のポリペプチドの変異体、配列番号4のポリペプチド、配列番号4と95%以上、98%以上、または99%以上の配列相同性を有するポリペプチド、および配列番号4のポリペプチドの変異体;
前記配列番号2のポリペプチドまたはこれと95%以上、98%以上、または99%以上の配列相同性を有するポリペプチドを暗号化するポリヌクレオチド、配列番号4のポリペプチドまたはこれと95%以上、98%以上、または99%以上の配列相同性を有するポリペプチドを暗号化するポリヌクレオチド、および前記ポリペプチドの変異体を暗号化するポリヌクレオチド;
前記ポリヌクレオチドを含む組換えベクター;および
前記組換えベクターを含む組換え細胞
からなる群より選択された1種以上を含む、植物および/または藻類の除草剤に対する耐性付与および/または増進用組成物を提供する。
(1)配列番号2、配列番号4、またはこれらと95%以上、98%以上、または99%以上の配列相同性を有するアミノ酸配列を含むポリペプチド、
(2)前述のポリペプチド変異体、
(3)前記ポリペプチド(1)またはポリペプチド変異体(2)を暗号化するポリヌクレオチド、
(4)前記ポリヌクレオチドを含む組換えベクター、および
(5)前記組換えベクターを含む組換え細胞
からなる群より選択された1種以上を含む、植物および/または藻類の除草剤に対する耐性付与および/または増進用組成物を提供する。
サーモシネココッカスエロンガータスBP−1(Thermosynechococcus elongatus BP−1)およびシネココッカス属(Synechococcus sp.)JA−3−3AbのGenBankデータベースを用いてPPO遺伝子情報を獲得した。BT3 E.coliでより効率的な除草剤抵抗性スクリーニングのために各PPO遺伝子をコドン最適化(codon optimization)して合成した(GenScript)。これを、pACBBベクターにクローニングするために表1のプライマーを用いて、下記の条件でPPO遺伝子を増幅した。
前記準備されたCyPPO10およびCyPPO13の除草剤耐性をPPO遺伝子が欠乏した大腸菌を用いて試験した。
前記準備されたCyPPO10およびCyPPO13遺伝子をpACBBベクター(Plasmid#32551;Addgene;図1参照)にクローニングした。
PCR産物30μl(microliter)、BamHIおよびXhoI(New England Biolabs)各0.5μl、10Xバッファー4μl、水(water)5.5μl;制限酵素反応(Restriction enzyme reaction)37℃、1時間
ライゲーション反応(Ligation reaction)は下記の条件下で行った:
T4 DNA ligase(RBC)0.5μl、Aバッファー1μl、Bバッファー1μl、前記の制限酵素で処理したPCR産物とベクター、総(total)10μl;22℃、30分
前記クローニングしたプラスミドをそれぞれBT3コンピテントセル(BT3 competent cell)に熱衝撃方法で形質転換した。各変異遺伝子種類別にクロラムフェニコール(Duchefa)が含まれているLB(Luria−Bertani)寒天培地で培養した。
PPO蛋白質と除草剤の結合構造情報の確認のために、PPO蛋白質の代表例としてCyPPO10を、PPO阻害除草剤の代表例としてチアフェナシル、サフルフェナシル、フルミオキサジン、スルフェントラゾンを使用して試験した。CyPPO10の蛋白質を暗号化する遺伝子をpET29bベクター(Catalog Number:69872−3;EMD Biosciences;図30参照)にクローニングし、大腸菌システムを使用してCyPPO10蛋白質を発現させた。発現されたCyPPO10蛋白質をニッケル親和クロマトグラフィーを通じて純粋分離し、精製されたCyPPO10蛋白質に各1mM濃度のチアフェナシル、サフルフェナシル、フルミオキサジン、またはスルフェントラゾンを添加してCyPPO10とチアフェナシル、サフルフェナシル、フルミオキサジン、またはスルフェントラゾンが結合した結晶を獲得した。その後、放射光加速器を用いて2.4Å解像度のCyPPO10とチアフェナシル、サフルフェナシル、フルミオキサジン、スルフェントラゾン結合体のX−線回折データを確保し、分子水準の3次元構造を糾明した。このような過程を通じて除草剤抵抗性を付与するCyPPO10蛋白質内アミノ酸変異位置に関する情報を収集した。
CyPPO10およびCyPPO13のPPO阻害除草剤耐性をより高めるためにそれぞれPPOアミノ酸配列中の前記実施例3で得られた除草剤と相互作用する位置のアミノ酸に変異を誘発してCyPPO10およびCyPPO13の変異遺伝子を製作した。
表3のプライマーを用いて、以下の条件下でPCRを行ってPPO遺伝子を分離および増幅させた:
PCR反応液条件
鋳型(CyPPO10またはCyPPO13の合成DNA) 1μl
10Xバッファー 5μl
dNTP混合物(各10mM) 1μl
フォワードプライマー(10μM) 1μl
リバースプライマー(10μM) 1μl
DDW 40μl
Pfu−X(Solgent、2.5unit/μl) 1μl
総(Total) 50μl
前記増幅した遺伝子産物(products)とpET303−CT Hisベクター(VT0163;Novagen;図3参照)をXbaI、XhoIで切断した後、T4 DNA ligase(RBC、3unit/μl)を用いてpET303−CyPPO10とpET303−CyPPO13プラスミドをそれぞれ製作した。
鋳型 1μ
10Xバッファー 5μl
dNTP混合物(各10mM) 1μl
フォワードプライマー(10μM) 1μl
リバースプライマー(10μM) 1μl
DDW 40μl
Pfu−X(Solgent、2.5unit/μl) 1μl
総(Total) 50μl
CyPPO10およびCyPPO13のPPO阻害除草剤耐性をより高めるためにそれぞれPPOアミノ酸配列中の前記実施例3で得られた除草剤と相互作用する位置のアミノ酸に変異を誘発した。このような変異を誘発させることができるPPO遺伝子を設計してPPOが欠乏したBT3大腸菌(BT3(ΔPPO))に形質転換した後、PPO阻害除草剤を処理した環境で培養して、形質転換大腸菌の生長阻害の有無を確認した。
前記実施例4で製作したpET303−CyPPO10とpET303−CyPPO13プラスミドとそれぞれの変異遺伝子が含まれているプラスミド(plasmid)をBT3コンピテントセル(BT3 competent cell)に熱衝撃方法で形質転換してアンピシリンが含まれているLB寒天培地で培養した。
前記得られた除草剤耐性をCyPPO野生型と比較した相対的水準で評価して下記の表8〜表11および図11〜図29に示した:
上記表8〜11で、野生型の除草剤耐性水準を「−」で表示し、これと同等水準の耐性を「−」で、高いほど「+」を付加して最大「+++++」まで等級を定めて評価した。
PPO蛋白質および特定位置のアミノ酸を変異させたPPO蛋白質変異体の酵素活性を測定しPPO阻害除草剤による阻害試験(inhibition assay)を行った。PPO蛋白質は水溶性が低いが、MBP(maltose binding protein)と共に融合タンパク質(fusion protein)(MBP−PPO)で発現させる場合、安定的に水溶性蛋白質が発現されることを確認し、下記の野生型および変異型蛋白質をMBPとの融合蛋白質形態に発現させて本試験に使用した(図4参照)。
誘導(Induction):OD600=0.2、最終濃度0.3mM IPTG添加;
発現温度:23℃、200rpmの振盪培養;
発現時間:16hrs
培養規模:200ml/1,000mlフラスコ(flask)。
抽出バッファー:カラムバッファー(Column buffer)(50mM Tris−Cl、pH8.0、200mM NaCl)5ml buffer/g cell;
超音波処理(Sonication):SONICS&MATERIALS社 VCX130(130watts);
15 sec ON、10 sec OFF for 5 min on ice;
4℃および20分条件下で遠心分離(20,000xg);および
前記遠心分離で得られた上澄み液をカラムバッファー(column buffer)を使用して1:6比率で希釈した。
前記結果から、CyPPO10とCyPPO13はシロイヌナズナPPO1(AtPPO1)、アマランサスPPO1よりVmax値が最少2.5倍から最大16倍以上高いのを確認することができた。結論的に、CyPPO10とCyPPO13のPPO蛋白質が植物由来のシロイヌナズナPPO01またはアマランサスPPO1よりPPO酵素としての能力がさらに優れるのを示す。
−チアフェナシル、サフルフェナシル、ホメサフェン、ブタフェナシル、フルミオキサジン、およびスルフェントラゾン各除草剤の最終濃度:0、10、50、100、250、500、1,000、2,500、5,000nM
IC50値は、前記酵素活性測定過程に除草剤を前記濃度で添加してPPO酵素活性を除草剤添加前の50%に阻害する除草剤の濃度として求めた。
上記表13から確認されるように、CyPPO蛋白質変異体の場合、野生型と比較して各除草剤のIC50値が顕著に上昇するのを確認することができる。このような結果は、PPO蛋白質の特定位置のアミノ酸変異によって除草剤に対する耐性が増加するのを立証するものである。本試験結果でCyPPO蛋白質変異体が大体的に野生型に比べて減少した酵素活性を有することが確認され、これは組換え蛋白質と植物で精製された蛋白質間の蛋白質折りたたみまたは疎水性が異なることに起因することと思われる。植物に存在するPPO蛋白質は植物体の葉緑体膜に存在し、疎水性であるが、大腸菌が生産する組換えPPO蛋白質はMBP蛋白質との融合蛋白質であって、親水性である。したがって、変異PPO蛋白質が植物の葉緑体で正常的な条件で発現されれば、変異蛋白質と野生型蛋白質間の酵素活性の差が大きくないことと思われる。
7−1.シロイヌナズナ形質転換ベクター製作およびシロイヌナズナ形質転換体製作
シロイヌナズナ形質転換は選別マーカであるbar遺伝子(グルホシネート耐性遺伝子)のORFとそれぞれのCyPPO10またはCyPPO13のアミノ酸変異体の暗号化遺伝子のORFを有するバイナリーベクター(binary vector)を製作して行った。PPO阻害除草剤と作用機序が異なる除草剤を交差使用する場合、その効果を確認するためにbar遺伝子を使用し、前記遺伝子は後代遺伝が安定的に行われているかどうかを検証することにも使用した。Bar遺伝子の発現のためにNOSプロモーターと転写終結のためのE9ターミネーターを使用した。
前記製作されたCyPPO10およびCyPPO13の野生型および変異遺伝子が挿入された形質転換シロイヌナズナの除草剤抵抗性を確認した。
図31a:25μMグルホシネート(PPT)、70nMチアフェナシル、100nMサフルフェナシル、25μMグルホシネート+70nMチアフェナシル、または25μMグルホシネート+30nMチアフェナシル+40nMサフルフェナシル;
図31bおよび31c:0.1μMまたは1μMチアフェナシル、0.3μMまたは3μMサフルフェナシル、0.1μMまたは1μMフルミオキサジン、0.5μMまたは5μMピラクロニル、または1μMまたは10μMスルフェントラゾン。
CyPPO10、CyPPO10変異体(F360I変異体またはF360M変異体)、CyPPO13、またはCyPPO13変異体(F373M変異体)を暗号化する遺伝子がそれぞれ挿入されたシロイヌナズナ形質転換体(T2)で各蛋白質が発現されるか調査した。
CyPPO10、CyPPO10変異体(F360C、F360I、F360L、F360M、F360V、F360T、A167C、A167L、A167L+F360M、A167C+F360M、A167C+F360I、またはV305M+F360M)、CyPPO13、またはCyPPO13変異体(A177C、F373C、F373I、F373M、A177L+F373I、またはA177L+F373L)を暗号化する遺伝子がそれぞれ挿入されたシロイヌナズナ形質転換体(T2またはT3)に対して除草剤抵抗性をテストした。
CyPPO10変異体(F360I、F360L、F360M、A167C+F360I、A167C+F360M、またはV305M+F360M)を暗号化する遺伝子がそれぞれ挿入されたシロイヌナズナ形質転換体(T3)にチアフェナシル溶液(25μMチアフェナシル+0.05%(v/v)シルウェットL−77)およびサフルフェナシル(100μMサフルフェナシル+0.05%(v/v)シルウェットL−77)を40×60cm面積(0.24m2)に各100mlずつ処理し、7日後に、植物の薬害程度を判断した。
チアフェナシル、サフルフェナシル、フルミオキサジン、スルフェントラゾン、オキシフルオルフェンまたはピラクロニル(各50μM)処理後7日目にCyPPO10変異遺伝子(F360I、またはA167L+F360M)またはCyPPO13変異遺伝子(A177L+F373L、またはA177L+F373I)が挿入されたシロイヌナズナ形質転換体(T3)の抵抗性水準を確認した。除草剤抵抗性と知られたシロイヌナズナPPO1の変異体(AtPPO1 SLYM、S305L+Y426M)が挿入されたシロイヌナズナ形質転換体(T3)またはシロイヌナズナ野生型(Col−0)を対照群として使用した。
図33aおよび表17で、「Cy10_FI」はCyPPO10 F360I変異遺伝子挿入個体、「AtPPO1_SLYM」はAtPPO1のS305L+Y426M変異遺伝子挿入個体(対照群)、「Cy10_ALFM」はCyPPO10 A167L+F360M変異遺伝子挿入個体での結果をそれぞれ示す。
本実施例では、シロイヌナズナ内挿入した遺伝子が世代を進展しても安定的に遺伝して発現されるか確認した。
形質転換体の世代別、ライン別に蛋白質を抽出した。具体的に、芽生え(Seedling)状態の植物体を液体窒素を用いて摩砕した後、蛋白質抽出バッファー(0.05M Tris−Cl pH7.5、0.1M NaCl、0.01M EDTA、1%トリトン(Triton)X−100、1mM DTT)を添加して総タンパク(total protein)を抽出し、これを用いてウエスタンブロットを行った。
約4週育った抽苔前シロイヌナズナ(T4およびT5CyPPO10 F360I形質転換体)を対象にして15μMチアフェナシルまたは150μMサフルフェナシルを40×60cm面積(0.24m2)に100mlずつ満遍なく撒布した。除草剤処理後7日目に除草剤薬害程度を観察した。
図34、図35、および表18に示されているように、対照群(Col−0;シロイヌナズナ野生型)は前記除草剤処理によって死滅する反面、T3で除草剤抵抗性を示した変異体(CyPPO10 F360I)のT4およびT5世代で除草剤抵抗性が維持されるのを確認することができる。これによって世代が進展しても変異体による除草剤抵抗性が安定的に伝達されるのを確認することができる。
8−1.大豆形質転換用組換えベクターおよびこれを用いた大豆形質転換体の製作
CyPPO10 A167L+F360M遺伝子を大豆植物体に発現させてチアフェナシル抵抗性を付与するために形質転換用ベクターを製作した。
CyPPO10 A167L+F360M形質転換大豆T0世代2番ラインおよび形質転換されていない大豆(クァンアン;野生型(wild type))の葉に5μMまたは15μMチアフェナシル溶液を筆を用いて2〜3回塗布した。チアフェナシル溶液は展着剤としてシルウェットL−77を0.05%(v/v)の濃度で含む。
CyPPO10A 167L+F360M形質転換大豆2、23番ラインの葉250mgでゲノムDNA(genomic DNA)抽出して形質転換大豆内の挿入遺伝子数を確認した。
1)脱プリン反応(depurination):0.25N HCl、15分間振盪(shaking)
2)変性(denaturation):0.5M NaOH、1.5M NaCl、30分間振盪(shaking)
3)中和(neutralization):0.5M Tris−Cl(pH7.5)、1.5M NaCl、20分間振盪(shaking)
そ の後、キャピラリートランスファー(capillary transfer)方法を用いてゲル内のDNA片をニトロセルロース膜(nitrocellulose membrane)(Amersham)に移した後、UV crosslinker(UVC−508;ULTRA LUM Inc.)を用いて架橋結合(cross linking)を行った。
Probe PCR
DIG dUTP(Jena bioscience)を用いてdigが標識されたbar遺伝子を増幅させ、この時使用されたプライマーは以下の通りである:
Forward primer for bar probe:5’−TTC CGT ACC GAG CCG CAG GA−3’(配列番号124)
Reverse primer for bar probe:5’−CGT TGG GCA GCC CGA TGA CA−3’(配列番号125)
PCR:Solgent e−Taqkit使用
条件:95℃5分後、94℃30秒、60℃30秒、72℃30秒を35cycle反復、72℃2分
前記反応した膜(membrane)に対してlow stringency washing(2X SSC、0.1%SDS)とhigh stringency washing(0.5X SSC、0.1%SDS)を行った。下記の条件でDIG検出(DIG detection)を行ってバンドを確認した:
1)膜をブロッキングバッファー(blocking buffer、Roche、11585762001)に入れて30分間振盪(shaking)
2)DIG抗体(anti−digoxigenin−AP Fab fragments、Roche)を入れて30分間振盪(shaking)
3)洗浄バッファー(Roche)で15分間振盪(shaking)
4)検出バッファー(Roche)を入れて3分間振盪(shaking)
5)CDP−Star、ready−to−use(Roche)を膜(membrane)に塗布した後、X線フィルム(x−ray film)でバンド(band)を現像する。
CyPPOプラスミド(pACBBベクター)を鋳型(template)にして以下の条件でエラープロンPCR(error−prone PCR)を行って、CyPPO内ランダム(random)突然変異を誘導した:
鋳型(Template) 0.5μl
10Xバッファー 5μl
10mM MnCl2 1.5μl
dNTP 5μl
e−Taq(Solgent社) 1μl
フォワードプライマー(forward primer)(100μM) 0.5μl
リバースプライマー(reverse primer)(100μM) 0.5μl
DDW 36μl
総50μl
10Xバッファー:100mM Tris−Cl、pH8.3;500mM KCl、70mM MgCl2、0.1%(w/v)ゼラチン(gelatin)
dNTP:10mM dATP、10mM dGTP、100mM dCTP、100mM dTTP
94℃ 3min;(94℃ 30sec、57℃ 30sec、72℃ 1.5min、72℃ 5min)35cycles
プライマー配列:
CyPPO10_BamHI F
ccccggatccATGATTGAAGTGGATGTGGCTA(配列番号126)
CyPPO10_XhoI R
ccccctcgagTGATTGTCCACCAGCGAGGTAAG(配列番号127)
CyPPO13_BamHI F
ccccggatccATGAACCCTGCTACCCCTGAAC(配列番号128)
CyPPO13_XhoI R
cccctcgagCACCTGTGATAACAACTGCTGAG(配列番号129)
前記得られたエラープロン(error−prone)PCR産物(PCR product)をアガロースゲル(agarose gel)に電気泳動した後、ゲル溶出(gel elution)し、pACBBベクターとPCR産物(PCR product)をBamHI、XhoI制限酵素で消化(digestion)した。制限酵素処理されたベクターとPCR産物(PCR product)をアガロースゲル(agarose gel)に電気泳動した後、ゲル溶出(gel elution)し、ライゲーション(ligation)した。ライゲーション産物(Ligation product)をBT3コンピテントセル(BT3 competent cell)に形質転換し、コロニーPCR(colony PCR)した後、突然変異CyPPOの配列(sequence)を確認した。突然変異が確認されたクローン(clone)をチアフェナシルまたはサフルフェナシルが濃度別(0、50、100、200μM)に含まれているLBプレート(LB plate)にスポッティング(spotting)して大腸菌の生長を調査した。突然変異クローンの中で、最終の下のクローンが除草剤抵抗性突然変異として選抜された:
CyPPO10m−6:9個アミノ酸変異(E225G、G258S、Q266L、T336I、V356F、F360M、A364D、R406G、W419R)含む;遺伝子配列−配列番号130、アミノ酸配列−配列番号131(野生型CyPPO10アミノ酸配列と98%配列相同性)
前記CyPPO10m−6の突然変異遺伝子を挿入したBT3形質転換体の除草剤抵抗性を確認して、その結果を図41に示した。図41で、「AtPPO1 WT」は野生型シロイヌナズナPPO1、「AtPPO1 SLYM」はY426MおよびS305L変異を含む変異型シロイヌナズナPPO1、「CyPPO10 WT」は野生型CyPPO10、「CyPPO10m−6」は前述のCyPPO10変異体をそれぞれ示す。
Claims (21)
- 配列番号2のアミノ酸配列中で、
(1)F360がM(Met)、V(Val)、I(Ile)、又はL(Leu)で置換され、R89がA(Ala)で置換され、V165がC(Cys)又はS(Ser)で置換され、A167がC(Cys)、L(Leu)、又はI(Ile)で置換され、V305がM(Met)で置換され、L327がT(Thr)で置換され、又はI408がR(Arg)又はW(Trp)で置換され;又は
(2)F360はM(Met)、V(Val)、I(Ile)、又はL(Leu)で置換され、そしてさらに
A(Ala)でのR89の置換;
C(Cys)又はS(Ser)でのV165の置換;
C(Cys)、L(Leu)、又はI(Ile)でのA167の置換;
M(Met)でのV305の置換;
T(Thr)でのL327の置換;及び
R(Arg)又はW(Trp)でのI408の置換
から選択される少なくとも1つの置換が導入されたアミノ酸配列を含む、ポリペプチド。 - 前記ポリペプチドは、
配列番号2のアミノ酸配列中で、
(1)F360がM(Met)、V(Val)、I(Ile)、又はL(Leu)で置換され;又は
(2)F360はM(Met)、V(Val)、I(Ile)、又はL(Leu)で置換され、そしてさらに
A(Ala)でのR89の置換;
C(Cys)又はS(Ser)でのV165の置換;
C(Cys)、L(Leu)、又はI(Ile)でのA167の置換;
M(Met)でのV305の置換;
T(Thr)でのL327の置換;及び
R(Arg)又はW(Trp)でのI408の置換
から選択される少なくとも1つの置換が導入されたアミノ酸配列を含む、請求項1に記載のポリペプチド。 - 前記ポリペプチドは、
配列番号2のアミノ酸配列において、F360M、F360V、F360I、F360L、A167C、A167L、R89A、V165S、V165C、A167I、V305M、L327T、I408R、I408W、R89A+F360M、R89A+F360I、R89A+F360L、R89A+A167L+F360M、V165S+F360M、V165S+F360I、V165S+F360L、V165S+F360V、V165C+F360M、V165C+A167C+F360M、V165C+A167I+F360M、V165C+A167L+F360M、A167L+F360M、A167L+F360I、A167C+F360M、A167C+F360I、A167I+F360M、V305M+F360M、I408R+F360M、I408W+F360M、L327T+F360M、R89A+F360V、V165C+F360V、V165C+F360I、V165C+F360L、A167C+F360V、A167C+F360L、A167I+F360V、A167I+F360I、A167I+F360L、A167L+F360V、A167L+F360L、V305M+F360V、V305M+F360I、V305M+F360L、V305L+F360V、L327T+F360V、L327T+F360I、L327T+F360L、I340T+F360M、I408R+F360I、I408R+F360V、I408W+F360I、I408W+F360L、I408W+F360V、R89A+V165C+F360M、R89A+V305M+F360M、R89A+L327T+F360M、V165C+V305M+F360M、V165S+L327T+F360M、V165C+F360V+I408R、V165C+F360V+I408W、A167C+V305M+F360M、A167L+F360V+I408R、A167L+F360V+I408W、V305M+F360V+I408R、V305M+F360V+I408W、R89A+V165C+F360I、R89A+V165S+F360M、R89A+V165S+F360I、R89A+A167C+F360M、R89A+A167C+F360I、R89A+A167L+F360I、R89A+V305M+F360I、R89A+I408R+F360M、R89A+I408W+F360I、V165C+A167C+F360I、V165C+A167C+F360V、V165C+A167I+F360I、V165C+A167I+F360L、V165C+A167I+F360V、V165C+A167L+F360I、V165C+A167L+F360L、A165C+A167L+F360V、V165C+V305M+F360I、V165C+V305M+F360L、V165C+V305M+F360V、V165C+I408R+F360M、V165C+I408R+F360I、V165C+I408R+F360L、V165C+I408R+F360V、V165C+I408W+F360M、V165C+I408W+F360I、V165C+I408W+F360L、V165C+I408W+F360V、A167C+V305M+F360L、A167C+V305M+F360V、A167L+V305M+F360M、A167L+V305M+F360I、A167L+V305M+F360L、A167L+V305M+F360V、A167L+I408R+F360M、A167L+I408R+F360I、A167L+I408R+F360L、A167L+I408W+F360M、A167L+I408W+F360I、A167L+I408W+F360L、V305M+I408R+F360M、V305M+I408R+F360I、V305M+I408R+F360L、V305M+I408W+F360M、V305M+I408W+F360I、V305M+I408W+F360L、R89A+V165S+A167C+F360M、R89A+V165S+V305M+F360M、R89A+V165S+V305M+F360L、R89A+V165S+L327T+F360M、R89A+A167L+V305M+F360M、R89A+A167L+V305M+F360V、V165S+A167C+V305M+F360M、V165S+A167C+L327T+F360M、R89A+V165S+A167C+F360L、R89A+V165S+A167C+F360V、R89A+V165S+V305M+F360I、R89A+A167L+V305M+F360I、V165C+A167L+V305M+F360M、V165C+A167L+V305M+F360I、V165C+A167L+V305M+F360L、V165C+A167L+V305M+F360V、V165S+A167L+V305M+F360I、V165C+A167L+I408R+F360M、V165C+A167L+I408R+F360L、V165C+A167L+I408W+F360M、V165C+A167L+I408W+F360I、V165C+A167L+I408W+F360L、V165C+V305M+I408R+F360M、V165C+V305M+I408R+F360I、V165C+V305M+I408R+F360L、V165C+V305M+I408W+F360M、V165C+V305M+I408W+F360I、V165C+V305M+I408W+F360L、A167L+V305M+I408R+F360M、A167L+V305M+I408R+F360I、A167L+V305M+I408R+F360L、A167L+V305M+I408W+F360M、A167L+V305M+I408W+F360I、A167L+V305M+I408W+F360L、R89A+V165S+A167C+V305M+F360M、R89A+V165S+V167C+V305M+F360I、R89A+V165S+A167C+L327T+F360M、R89A+V165S+V305M+L327T+F360M、V165S+A167C+V305M+L327T+F360M、R89A+V165C+A167L+V305M+F360I、V165C+A167L+V304M+I408R+F360M、V165C+A167L+V305M+I408R+F360I、V165C+A167L+V305M+I408R+F360L、V165C+A167L+V305M+I408W+F360M、V165C+A167L+V305M+I408W+F360I、又はV165C+A167L+V305M+I408W+F360Lのアミノ酸変異を含むアミノ酸配列を含む、請求項1に記載のポリペプチド。 - 請求項1〜3のいずれか一項のポリペプチドをコードするポリヌクレオチド。
- 請求項4のポリヌクレオチドを含む組換えベクター。
- 請求項5の組換えベクターを含む組換え細胞。
- 請求項1〜3のいずれか一項のポリペプチド;前記ポリペプチドをコードするポリヌクレオチド;前記ポリヌクレオチドを含む組換えベクター;および前記組換えベクターを含む組換え細胞からなる群より選択される少なくとも1つを含む、植物または藻類(algae)の除草剤に対する抵抗性付与または増進のための組成物であって、除草剤はプロトポルフィリノーゲンオキシダーゼを抑制する除草剤である、組成物。
- 前記除草剤は、ピリミジンジオン系、ジフェニルエーテル系、フェニルピラゾール系、N−フェニルフタルイミド系、フェニルエステル系、チアジアゾール系、オキサジアゾール系、トリアゾリノン系、オキサゾリジンジオン系、ピラクロニル、フルフェンピル−エチルおよびプロフルアゾルからなる群より選択される少なくとも1つである、請求項7に記載の組成物。
- 前記除草剤は、ブタフェナシル、サフルフェナシル、ベンズフェンジゾン、チアフェナシル、ホメサフェン、オキシフルオルフェン、アクロニフェン、アシフルオルフェン、ビフェノックス、エトキシフェン、ラクトフェン、クロメトキシフェン、クロルニトロフェン、フルオログリコフェン−エチル、ハロサフェン、ピラフルフェン−エチル、フルアゾレート、フルミオキサジン、シニドン−エチル、フルミクロラック−ペンチル、フルチアセット、チジアジミン、オキサジアルギル、オキサジアゾン、カルフェントラゾン、スルフェントラゾン、アザフェニジン、ペントキサゾン、ピラクロニル、フルフェンピル−エチル、プロフルアゾル、フェノピレート、フェノピレートのカルバメート類似体、これらの農薬的に許容可能な塩からなる群より選択される少なくとも1つである、請求項7又は8に記載の組成物。
- 上記植物または藻類は、第2の除草剤抵抗性ポリペプチドまたはこれをコードする遺伝子をさらに含み、前記第2の除草剤に対する抵抗性が付与または増進される、請求項7〜9のいずれか一項に記載の組成物。
- 前記第2の除草剤は、グリホサート、グルホシネート、ジカンバ、2,4−D(2,4−ジクロロフェノキシ酢酸)、イソキサフルトール、ALS(アセト乳酸シンターゼ)阻害性除草剤、光化学系II阻害性除草剤、フェニルウレア系除草剤、ブロモキシニル系除草剤およびこれらの組み合わせからなる群より選択される、請求項10に記載の組成物。
- 前記第2の除草剤耐性ポリペプチドは、
グリホサート除草剤耐性EPSPS(glyphosate tolerant 5−enolpyruvylshikimate−3−phosphate synthase)、GOX(グリホサートオキシダーゼ)、GAT(グリホサート−N−アセチルトランスフェラーゼ)またはグリホサートデカルボキシラーゼ;
グルホシネート除草剤耐性PAT(ホスフィノトリシン−N−アセチルトランスフェラーゼ);
ジカンバ除草剤耐性DMO(ジカンバモノオキシゲナーゼ);
2,4−D除草剤耐性2,4−DモノオキシゲナーゼまたはAAD(アリールオキシアルカノアートジオキシゲナーゼ);
ALS阻害性スルホニルウレア系除草剤耐性ALS(acetolactate synthase)、AHASまたはAtAHASL;
光化学系II阻害性除草剤耐性光化学系IIタンパク質D1;
フェニルウレア除草剤耐性シトクロムP450;
色素体阻害性除草剤耐性HPPD;
ブロモキシニル除草剤耐性ニトリラーゼ;および
これらの組み合わせからなる群より選択される少なくとも1つである、請求項10に記載の組成物。 - 前記第2の除草剤耐性ポリペプチドをコードする遺伝子は、
グリホサート除草剤耐性cp4 epsps、epsps(AG)、mepsps、2mepsps、goxv247、gat4601またはgat4621遺伝子;
グルホシネート除草剤耐性barまたはpat遺伝子;
ジカンバ除草剤耐性dmo遺伝子;
2,4−D除草剤耐性AAD−1またはAAD−12遺伝子;
イソキサフルトール除草剤耐性HPPDPF W336遺伝子;
スルホニルウレア除草剤耐性ALS、Csr1、Csr1−1、Csr1−2、GM−HRA、S4−HRA、Zm−HRA、SurAまたはSurB遺伝子;
光化学系II阻害性除草剤耐性psbA遺伝子;
フェニルウレア除草剤耐性CYP76B1遺伝子;
ブロモキシニル除草剤耐性bxn遺伝子;および
これらの組み合わせからなる群より選択される少なくとも1つである、請求項10に記載の組成物。 - 請求項1〜3のいずれか一項のポリペプチドまたはそれらをコードするポリヌクレオチドを含む、植物または藻類の除草剤に対する耐性を有する形質転換体、そのクローンまたは子孫であって、除草剤はプロトポルフィリノーゲンオキシダーゼを抑制する阻害剤である、形質転換体、そのクローンまたは子孫。
- 前記形質転換体は、植物の細胞、プロトプラスト、カルス、胚軸、種子、子葉、シュートまたは植物体全体である、請求項14に記載の形質転換体、そのクローンまたは子孫。
- 請求項1〜3のいずれか一項のポリペプチドをコードするポリヌクレオチドを藻類、または植物の細胞、プロトプラスト、カルス、胚軸、種子、子葉、シュートまたは植物体全体に形質転換することを含む、除草剤に対する耐性を有する植物または藻類の製造方法であって、除草剤はプロトポルフィリノーゲンオキシダーゼを抑制する阻害剤である、製造方法。
- 請求項1〜3のいずれか一項のポリペプチドをコードするポリヌクレオチドを藻類、または植物細胞、プロトプラスト、カルス、胚軸、種子、子葉、シュートまたは植物体全体に形質転換することを含む、植物または藻類の除草剤に対する耐性を付与または増進させる方法であって、除草剤はプロポルフィリノーゲンオキシダーゼを抑制する除草剤である、方法。
- 請求項1〜3のいずれか一項のポリペプチド、またはそれらをコードするポリヌクレオチドを含む植物を栽培地に提供すること、および
前記栽培地にプロトポルフィリノーゲンオキシダーゼを抑制する除草剤の有効量を適用することを含む、
栽培地で雑草を防除する方法。 - 前記栽培地にプロトポルフィリノーゲンオキシダーゼを抑制する除草剤の有効量を適用することは、2つ以上のプロトポルフィリノーゲンオキシダーゼを抑制する除草剤の有効量を順次にまたは同時に適用することである、請求項18に記載の方法。
- 上記植物は、第2の除草剤耐性ポリペプチドまたはこれをコードする遺伝子をさらに含み、
前記栽培地にプロトポルフィリノーゲンオキシダーゼを抑制する除草剤の有効量を適用することが、プロトポルフィリノーゲンオキシダーゼを抑制する除草剤および第2の除草剤の有効量を順次にまたは同時に適用することにより実施される、請求項18に記載の方法。 - 請求項1〜3のいずれか一項のポリペプチド、またはそれらをコードするポリヌクレオチドを含む藻類を培養培地に提供すること、および
前記培養培地にプロトポルフィリノーゲンオキシダーゼを抑制する除草剤の有効量を適用することを含む、
培養培地で望まない水生生物を除去する方法。
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CL2018003620A1 (es) | 2019-03-29 |
AR108807A1 (es) | 2018-09-26 |
UY37298A (es) | 2018-01-02 |
EP3472184A4 (en) | 2019-11-20 |
US20210095305A1 (en) | 2021-04-01 |
IL263413B2 (en) | 2024-02-01 |
KR101862796B1 (ko) | 2018-05-30 |
CN109476709A (zh) | 2019-03-15 |
MX2018015794A (es) | 2019-03-21 |
KR102250017B1 (ko) | 2021-05-10 |
US10844395B2 (en) | 2020-11-24 |
AU2017284680A1 (en) | 2018-12-13 |
ZA201807594B (en) | 2020-02-26 |
IL263413B1 (en) | 2023-10-01 |
KR20190127644A (ko) | 2019-11-13 |
KR102044430B1 (ko) | 2019-11-14 |
AU2017284680B2 (en) | 2020-04-30 |
WO2017217793A1 (en) | 2017-12-21 |
US20190330651A1 (en) | 2019-10-31 |
CL2020001807A1 (es) | 2020-12-04 |
CA3024985A1 (en) | 2017-12-21 |
PH12018502621A1 (en) | 2019-10-07 |
IL263413A (en) | 2018-12-31 |
JP2019527538A (ja) | 2019-10-03 |
EP3472184A1 (en) | 2019-04-24 |
US11466286B2 (en) | 2022-10-11 |
KR20170142120A (ko) | 2017-12-27 |
KR20180000717A (ko) | 2018-01-03 |
CN116064434A (zh) | 2023-05-05 |
CO2019000095A2 (es) | 2019-02-19 |
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