JP7399375B2 - 接木改善剤 - Google Patents
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- JP7399375B2 JP7399375B2 JP2019052727A JP2019052727A JP7399375B2 JP 7399375 B2 JP7399375 B2 JP 7399375B2 JP 2019052727 A JP2019052727 A JP 2019052727A JP 2019052727 A JP2019052727 A JP 2019052727A JP 7399375 B2 JP7399375 B2 JP 7399375B2
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Description
(B)配列番号1若しくは2に示されるアミノ酸配列、又は配列番号1若しくは2に示されるアミノ酸配列に対して1又は数個のアミノ酸が置換、欠失、付加若しくは挿入されてなるアミノ酸配列を含み、且つ接木改善活性を有するペプチド、
(C)フェノールアミド類又はポリアミン、及び
(D)前記(A)~(C)のいずれかの発現又は生成促進剤
からなる群より選択される少なくとも1種を含有する、接木改善剤。
前記(B)の発現及又は生成促進剤が前記(B)の発現カセットであり、
前記(C)の発現及又は生成促進剤が前記(C)の生合成関連遺伝子の発現カセット又は前記(C)の生合成関連因子である、
項1又は2に記載の接木改善剤。
本明細書中において、「含有」及び「含む」なる表現については、「含有」、「含む」、「実質的にからなる」及び「のみからなる」という概念を含む。
担体としては、本発明のポリペプチドを担持可能なものである限り特に制限されない。担体の材質としては、特に制限されず、各種樹脂等の有機材料、ケイ素材料や金属等の無機材料等が例示される。
本発明は、その一態様において、(A)セルラーゼ、(B)配列番号1若しくは2に示されるアミノ酸配列、又は配列番号1若しくは2に示されるアミノ酸配列に対して1又は数個のアミノ酸が置換、欠失、付加若しくは挿入されてなるアミノ酸配列を含み、且つ接木改善活性を有するペプチド、(C)フェノールアミド類又はポリアミン、及び(D)前記(A)~(C)のいずれかの発現又は生成促進剤からなる群より選択される少なくとも1種を含有する、接木改善剤(本明細書において、「本発明の接木改善剤」と示すこともある。)に関する。以下に、これについて説明する。なお、(A)、(B)、(C)、(D)の成分については、それぞれ、A成分、B成分、C成分、D成分と示すこともある。
セルラーゼとしては、特に制限されず、各種セルラーゼを採用することができる。セルラーゼとしては、例えばエンドグルカナーゼ(EC3.2.1.4)、エキソグルカナーゼ(EC.3.2.1.91)が挙げられる。セルラーゼとしては、ファミリー内の各種セルラーゼを使用することができ、例えばグリコシルヒドロラーゼファミリーが挙げられ、具体的には、例えばグリコシルヒドロラーゼ1、グリコシルヒドロラーゼ2、グリコシルヒドロラーゼ3、グリコシルヒドロラーゼ4、グリコシルヒドロラーゼ5、グリコシルヒドロラーゼ6、グリコシルヒドロラーゼ7、グリコシルヒドロラーゼ8、グリコシルヒドロラーゼ9、グリコシルヒドロラーゼ10、グリコシルヒドロラーゼ11、グリコシルヒドロラーゼ12等が挙げられる。これらの中でも、特に好ましくはグリコシルヒドロラーゼ9(例えば9A、9B、9Cなど、中でも好ましくは9B、より好ましくは9B1、9B13)等が挙げられる。
(i)配列番号3又は4に示されるアミノ酸配列からなるタンパク質、又は
(ii)配列番号3又は4に示されるアミノ酸配列と70%以上の同一性を有するアミノ酸配列からなり、且つ接木改善活性を有するを有するタンパク質
である。
(iia)配列番号3又は4に示されるアミノ酸配列に対して1若しくは複数個のアミノ酸が変異(例えば置換、欠失、付加、挿入等、好ましくは置換、より好ましくは保存的置換)したアミノ酸配列からなり、且つ接木改善活性を有するタンパク質
である。
B成分は、配列番号1若しくは2に示されるアミノ酸配列、又は配列番号1若しくは2に示されるアミノ酸配列に対して1又は数個のアミノ酸が置換、欠失、付加若しくは挿入されてなるアミノ酸配列を含み、且つ接木改善活性を有するペプチドである。
本発明において、C成分としての「フェノールアミド類」とは、フェノール(但し2以上のヒドロキシ基を有していても、またヒドロキシ基以外の置換基(例えばアルコキシ基)を1以上有していてもよい。)を含むアシル基と、(ポリ)アミン残基とが、1以上のアミド結合で連結した化合物群を意味する。植物内では、フェニルプロパノイド類のアシル基が、ポリアミンに転移(結合)して、フェノールアミド(別称として、hydroxycinnamic acid amide (HCAA)ともいう)類が産生する現象が知られている。本発明に利用されるフェノールアミド類には、この様な植物内に存在する天然のフェノールアミド類が含まれる。
A成分の発現又は生成促進剤としては、A成分の発現又は生成を促進可能なものである限り特に制限されず、例えばA成分(セルラーゼ)の発現カセット、A成分(セルラーゼ)の遺伝子の転写活性化因子の発現カセット、A成分(セルラーゼ)の遺伝子の転写抑制因子の発現又は機能阻害材(例えば抗体、siRNA発現カセット、CRISPR/Casシステム発現カセット等)等が挙げられる。
本発明の接木改善剤は、A成分、B成分、C成分、及びDS成分からなる群より選択される少なくとも1種を有効成分として含有する。本発明の接木改善剤は、より具体的には、例えば、接木効率向上のために、接木成功率向上のために、接木和合性向上のために、接木成立速度向上のために、接木失敗率低減のために、使用することができる。
本発明は、その一態様において、植物体又は植物部分構造体に本発明の接木接着剤を施用することを含む、接木用植物部分構造体の製造方法、及び該方法により得られた接木用植物部分構造体に関する。
セルラーゼ遺伝子をノックダウンしたタバコ属植物(Nicotiana benthamiana)(穂木)とシロイヌナズナ(Arabidopsis thaliana)(台木)とを接木し、接木成立の程度を評価した。具体的には以下のようにして行った。
試験例1と同様にタバコ属植物とシロイヌナズナとの接木試験を行い、壊死層領域の頻度を調べた。具体的には以下のようにして行った。
セルラーゼ投与による接木成立性への影響を、In vitro graftingの試験系により評価した。具体的には以下のようにして行った。
配列番号1のアミノ酸配列(MVTDHVKNAT)からなるペプチド(Grf006)と配列番号2のアミノ酸配列(MALIVQMISA)からなるペプチド(Grf010)の接木改善作用を、In vitro graftingの試験系により評価した。具体的には以下のようにして行った。
タバコ属植物(Nicotiana attenuata)においては、NaMYB8がフェノールアミド生合成のマスターレギュレーターであることが知られている(Kaur et al., Plant Physiology, March 2010, Vol. 152, pp. 1731-1747、Onkokesung et al., Plant Physiology, January 2012, Vol. 158, pp. 389-407)。本試験では、トマトの野生株に対して、NaMYB8のオーソログであるSIMYB14を、RNAi法でノックダウンしてRNAi体(2-4-1)を製造し、この2-4-1体のSIMYB14遺伝子発現量、及び代謝物(フェノールアミド類)の蓄積量を、野生株と対比するとともに、接木への影響を評価した。
SIMYB14遺伝子の部分配列(323 bp)をRNAiのトリガー配列とし、ユビキチンプロモータ下でRNAiのトリガー配列を発現させる形質転換ベクター(pANDA)をアグロバクテリウム法により、トマト‘Ailsa Craig’の子葉に形質転換し、得られた形質転換シュートを発根させ、形質転換体(RNAi体(2-4-1))を得た。形質転換体作出過程で得られた非形質転換個体(すなわち野生型トマトと同形質の個体)をEscape個体(2-1-1体)として以下の実験に用いた。
野生株、並びに上記で製造した2-4-1体及び2-1-1体それぞれからTotal RNAを抽出し、逆転写反応を行い、cDNAを得た。得られたcDNAを鋳型として、SIMYB14の部分長を増幅するプライマーを用い、定量PCR法により、SIMYB14遺伝子発現量を測定した。
SlMYB14RNAi体(2-4-4)の凍結粉末サンプルとジルコニアビーズ1個が入ったチューブに1 mg新鮮重当たり5 μlの80%メタノール(内部標準として2.5 μMのリドカインを含む)を加え、ミキサーミルで攪拌抽出を行った。当該チューブを遠心分離にかけ、残渣を沈殿させ、上清を新しいチューブに移すことで抽出液を得た。
液体クロマトグラフィー(LC)部: カラム, Acquity bridged ethyl hybrid C18 (1.7 μm, 2.1 mm × 100 mm, Waters);溶媒系:溶媒 A (0.1% ギ酸を含む水) 及び 溶媒 B (0.1% ギ酸を含むアセトニトリル);濃度勾配: 99.5% 溶媒A/0.5% 溶媒B、0分まで、99.5%A/0.5%Bを0.1分まで、20%A/80%Bを10分まで、0.5%A/99.5%Bを10.1分まで、0.5%A/99.5%Bを12.0分まで、99.5%A/0.5%Bを12.1分まで、99.5%A/0.5%Bを15.0 分まで;流速:0.3 ml/minを0 minまで, 0.3 ml/minを10分まで、0.3 ml/minを10.1分まで、0.3 ml/mlを14.4分まで、0.3 ml/minを14.5分まで:カラム内温度: 40 度;質量分析(MS)部:極性, ポジティブおよびネガティブイオンモード;キャピラリーボルテージ、+3.00 kV (ポジティブ)/-2.75 kV (ネガティブ);コーンボルテージ: 25.0 V;イオンソース内温度:120 ℃;デソルベーション温度:450 ℃;コーン内ガス流速: 50 l/h;デソルベーションガス流速: 800 l/h;コリジョンエナジー:6 V;質量分析範囲:m/z 50-1,500;スキャンデュレーション:0.1 s;inter-scan delay: 0.014 s;データ取得方法:セントロイドモード;ロックスプレー (ロイシンエンケファリン);スキャンデュレーション:1.0 s; inter-scan delay:0.1 s.。
(1) MS:極性, ポジティブおよびネガティブイオンモード;質量分析範囲, m/z50-1,500;スキャンデュレーション、0.1 s;interscan delay、0.014 s;データ取得方法:セントロイドモード。
(2) MS/MS:極性, ポジティブおよびネガティブイオンモード;質量分析範囲, m/z50-1,500;スキャンデュレーション、0.02 s; inter-scan delay、0.014 sデータ取得方法:セントロイドモード。
次に、フェノールアミドのナス科植物における普遍性を、種々の品種のナス科植物をサンプルとして評価した。具体的には以下の通りである。
N. benthamiana(シロイヌナズナ)とA. thaliana(タバコ)の接ぎ木植物、並びにN. benthamiana(シロイヌナズナ)の傷処理穂木及び台木をそれぞれ準備した。接ぎ木植物については、接木処理から28日間育成し、2時間後、及び1日後、3日後、5日後、7日後、10日後、14日後及び28日後に、それぞれ試料を採取して、上記と同様にして、メタボローム解析を行い、フェノールアミド類の構造決定及び定量を行った。また、シロイヌナズナの傷処理穂木及び台木についても、7日間育成し、2時間後、及び1日後、3日後、5日後及び7日後に、それぞれ試料を採取して、上記と同様にして、メタボローム解析を行い、フェノールアミド類の構造決定及び定量を行った。それぞれの結果を、なんら処理をしていないシロイヌナズナ及びタバコからそれぞれ採取した試料(参照例)のメタボローム解析結果と対比したところ、参照例と比較して、明らかに、フェノールアミド類の量が時間経過で変動していることが確認された。
野生株、並びに上記で製造した2-4-1体及び2-1-1体それぞれについて、セルフ接ぎ木(切り取った茎を元の茎に接ぎ木)し、活着(穂木の成長)を評価した。
フェノールアミド類生合成経路上の中間体(ポリアミン)の接木改善作用を、In vitro graftingの試験系により評価した。具体的には以下のようにして行った。
Claims (6)
- (A)セルラーゼ、
及び
(D)プロモーター及びそのプロモーターの制御下に配置されたセルラーゼコード配列を含むポリヌクレオチドからなる発現カセット
からなる群より選択される少なくとも1種を含有する、接木改善剤。 - 異科接木用又は同科接木用である、請求項1に記載の接木改善剤。
- 植物体又は植物部分構造体に請求項1又は2に記載の接木改善剤を施用することを含む、接木用植物部分構造体の製造方法。
- 請求項3に記載の方法により得られた、接木用植物部分構造体。
- 接木後の植物体に請求項1又は2に記載の接木改善剤を施用すること、及び/又は請求項4に記載の接木用植物部分構造体を他の植物部分構造体と接木することを含む、接木植物体の製造方法。
- 請求項5に記載の方法により得られた、接木植物体。
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