JP2952041B2 - 培養ダイズ細胞のａｇｒｏｂａｃｔｅｒｉｕｍ媒介形質転換の改良法 - Google Patents

培養ダイズ細胞のａｇｒｏｂａｃｔｅｒｉｕｍ媒介形質転換の改良法

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JP2952041B2
JP2952041B2 JP6504746A JP50474693A JP2952041B2 JP 2952041 B2 JP2952041 B2 JP 2952041B2 JP 6504746 A JP6504746 A JP 6504746A JP 50474693 A JP50474693 A JP 50474693A JP 2952041 B2 JP2952041 B2 JP 2952041B2
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エイ． タウンセンド，ジェフリー
エイ． トーマス，ローリー
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Description

【発明の詳細な説明】 発明の分野 本発明は、Agrobacterium媒介形質転換によるトラン
スジェニックダイズ植物の生成に関する。
発明の背景 ダイズ（Glycine max）は、世界で最も重要な作物植
物の１つである。油およびタンパク質の生産のために、
5000万ヘクタールを超えて生育されている１年生マメ科
植物である。作物の生産価値は、200億ドルを越えると
評価される。毎年、１億メートルトンを超えるダイズが
生産される。マメ科植物の遺伝子転移技術の開発は、耐
病性、除草剤に対する抵抗性、および栄養価を増大する
ように改良した新規な栽培品種の開発が容易となるた
め、商業的に重要である。しかし、ダイズの改良のため
の分子学的アプローチは、トランスジェニックダイズ植
物を生成するのに利用可能な技術に制限される。
いくつかの重要な作物植物を包含する多くの植物は、
Agrobacterium媒介遺伝子転移を用いて遺伝的に改変さ
れている。McCormickら，（1986）Plant Cell Rep.5,81
−84;Radkeら，（1988）Theor.Appl.Genet.75,685−69
4;Umbeckら，（1987）Bio/Technology 5,263−266;Bott
emanおよびLeemans,（1988）Trend in Genetics 4,219
−222。不運にも、これらの双子葉植物種のいくつかの
遺伝子型はAgrobacterium感染に感受性であっても（Fac
ciottiら，（1985）Biotechnology 3,241−246;Owensお
よびCress（1985）Plant Physiol.77,87−94;Byrneら，
（1987）Plant Cell Tissue and Organ Culture 8,3−1
5）、形質転換にAgrobacteriumを使用することは、利用
可能で有効な形質転換および再生の手順がないことによ
り制限される。
最近まで、ダイズはAgrobacteriumの宿主範囲外にあ
ると考えられていた。DeCleeneおよびDeLey（1976）Bac
terial Rev.42,389−466。最近の報告では、ダイズは、
ダイズ遺伝子型およびAgrobacterium株に依存して、Agr
obacteriumに対して感受性が制限されていることを示唆
した。Byrneら，（1987）Plant Cell Tissue Organ Cul
t.8,3−15。たった１種のダイズのみ（Peking）が形質
転換に成功しているが、この変種は市場価値がない。こ
の形質転換可能な変種のわずかなトランスジェニックダ
イズ植物がAgrobacteriumと再生可能な外植片との共存
培養により生成されている。Hincheeら，（1988）Bio/T
echnology 6,915−922。Agrobacterium媒介法によるダ
イズの形質転換が市販可能になるためには、この方法
は、より抜きの商業的栽培品種の直接的形質転換が可能
になるようにしなければならない。
これらの問題に加えて、ダイズは再生が最も困難なGl
ycine種であることが知られている。外植片は根を容易
に生成せず、さらにトランスジェニック材料の栽培を行
うことはほとんど不可能である。このため、形質転換お
よび再生に用いられる方法は、現在まで信頼性がなく効
果がないままであった。
従って、本発明の目的は、植物細胞に感染し、その植
物細胞にＴ−DNAを転移してそこで発現させる、Agrobac
terium種の改良法を提供することである。
本発明の別の目的は、ダイズ植物細胞に感染し、この
細胞を形質転換するAgrobacterium種の能力を高めるた
めの、この種の新規な誘導培地を提供することである。
本発明のさらに別の目的は、ダイズ外植片から完全植
物への再生のための、新規な成長培地および方法を提供
することである。
本発明のプロセスは、ダイズ組織でのAgrobacterium
媒介形質転換技術およびそのトランスジェニックダイズ
植物への再生において大きな改良を示す。
発明の要旨 本発明に従って、トランスジェニックダイズ植物の生
成のための簡単で迅速な信頼性のあるプロセスが提供さ
れる。方法は、より抜きの商業的栽培品種を包含するす
べてのダイズ変種に有効であり、今までに用いられたシ
ステム以上に相当な改良を示す。これは、この方法が、
ダイズの形質転換および再生に必要でありこれまで欠け
ていた要因を提供し、そして健常で可稔性のトランスジ
ェニック植物生成を成功させるために、これらおよび他
の要因を最適化するからである。
本発明は、以下の工程を包含する、遺伝子型に依存し
ないトランスジェニックダイズ植物の生成方法を包含す
る： （ａ）発芽させたダイズ種子の下胚軸または培養子葉節
由来の外植片を、キメラ遺伝子を含むAgrobacterium種
と共に、アセトシリンゴン、α−ヒドロキシアセトシリ
ンゴン、アセトバニロン、シリンガアルデヒド、シリン
ガ酸、およびシナピン酸、ならびにそれらの混合物から
なる群から選択されるシグナル化合物の存在下で共存培
養する工程； （ｂ）28℃より低い温度での共存培養によりAgrobacter
iumの毒性を誘導する工程；および （ｃ）植物培養培地のpHをpH6.0より低く低下させるこ
とによりAgrobacteriumの毒性を誘導する工程。
このプロセスは、ダイズ下胚軸または子葉節外植片
と、目的の１つまたは複数の遺伝子を挿入されるプラス
ミドを有するAgrobacterium種との共存培養を包含す
る。ダイズ外植片は、供給植物から採取され、培養にお
いてカルスを生成し得る一片のダイズ組織である。ダイ
ズ下胚軸組織は、子葉の下部で根の上部のダイズ植物胚
または実生の軸の部分である。子葉は胚葉であり、そし
て子葉節は胚軸と子葉との間の実生部であり、これは下
胚軸と上胚軸との区別を植物学的に規定する（胚シュー
ト）。
培養ダイズ細胞の形質転換にかなり影響するいくつか
の要因が、本発明の達成において同定された。これらの
うち最も重要なことは、共存培養の間シグナル分子の正
しい使用によるAgrobacteriumの毒性（vir）遺伝子の誘
導であると思われる。培養ダイズ細胞は、形質転換プロ
セスを開始させるのに必要なシグナル分子を有さない
か、または限られた量で有する。これらの結果は、一般
に、ダイズ形質転換のためのvir遺伝子の導入の重要性
を認めているがこの問題を解決していない他の研究と一
致する。Delzerら，（1990）Crop Sci.30,320−322;Owe
nsおよびSmigocki（1988）Plant Physiol.88,570−57
3。本発明は、アセトシリンゴン（傷害を受けた植物細
胞により生成されるフェノール性化合物）を用いて、vi
r遺伝子を誘導する。他のフェノール性化合物、α−ヒ
ドロキシアセトシリンゴン、アセトバニロン、シリンガ
アルデヒド、シリンガ酸、およびシナピン酸もまたvir
遺伝子を誘導し、独立してまたはアセトシリンゴンと組
み合わせたそれらの使用により、ダイズ形質転換の効果
が改良され得る。共存培養プロセスで十分な量のシグナ
ル分子を使用することにより、あらゆる場合において、
形質転換頻度が高められた。
共存培養の温度が別の重要な要因であることが見出さ
れた。ダイズ細胞培養の通常の温度（26〜28℃）は、有
効な形質転換には不適切である。温度をより低くするこ
とにより、より有効な形質転換が行われる。28℃で観察
された阻害は、より高い温度の最近の過増殖およびその
結果生じる植物細胞の生存の低下から生じるようではな
かった。これは、硫酸カナマイシンを培地から除去した
とき、両温度で共存培養した外植片は大きなカルスを生
成したからである。共存培養中の培地pHもまた、形質転
換率に影響する。低pH（pH 5.5）に緩衝化するような処
理により高い効果が確実となる。理論により制限される
ことを意図しないが、これらの改良効果はシグナル分子
反応作用に関連すると思われる。vir遺伝子誘導は温度
およびpHに依存し、最適条件は約20℃およびpH 5.5であ
ることが知られている。Alt−Moerleら，（1988）Mol.G
en.Genet.213,1−８。
ダイズ細胞の形質転換の成功はまた、接種物における
細菌の濃度にも依存する。一般に、細菌数が大きくなる
ことにより、より多くの形質転換が達成される。飽和濃
度の細菌が存在し得るが、ここではダイズ細胞ではさら
なる形質転換は生じず、この数は非常に大きいと思われ
る。本発明者らの実験は、この数が、３×108生存細胞/
mlで、あるいはそれを超える濃度で細菌を１回30分接種
することにより行い得、そして３×107細胞/mlの従来濃
度（Deblaereら，（1987）Meth.Enzymology 153,277−2
93）では、かなり低すぎることを示唆した。
しかし、本発明者らの実験では、真の接種飽和条件は
特定され得なかった。連続接種により形質転換可能なダ
イズ細胞は、接種が非常に高い細菌濃度で行われるとき
であっても標的されることがないことが示された。高濃
度の細菌を接種しようと試みるときに直面する問題の１
つは、細菌細胞の凝集の傾向であった。凝集により、植
物細胞に付着するのに用いられ得る細菌数が減少した。
沈降した細菌を再懸濁するとすぐに凝集が始まり、この
凝集率は濃度依存性であった。これはまた、培地の機械
的混合の程度にも依存した。共存培養培地（細菌が再懸
濁される）の組成もまた凝集に影響し得る。従って、植
物細胞の急速な***の誘導およびAgrobacterium vir遺
伝子の高レベル誘導に適切な完全植物細胞培地のみがこ
のプロセスにおいて用いられた。
形質転換頻度の相当な増大は、凝集現象を制限し、そ
の効果による接種条件を用いて達成された。第一に、接
種は、混合を最小限にして、室温で高い開始濃度で行っ
た。第二に、それらは、各外植片に対し新しく再懸濁し
た対数増殖期のAgrobacteriumのペレットを接種するよ
うに、バッチ式で行った。外植片は、可能な限り最も高
い濃度での利用可能な細菌の存在下で（すなわち、個々
の細胞数をかなり減少させる凝集をする前に）、接種物
で傷つけた。
第三に、接種期間は、所定のバッチの最後の外植片を
傷害してから開始して30分ぐらいとした。時間をこれよ
り長くしても形質転換頻度を増大することはなかった。
連続接種により、頻度を適度に増大させた。この増大
は、おそらく植物細胞を形質転換するのに利用可能な細
菌の数の増加によると思われた。これらの実験から、ダ
イズ細胞は、高濃度のAgrobacteriumに長期間さらされ
ることに耐え得ることが明らかである。
vir遺伝子誘導を確実にする条件を用い、必要な高濃
度のAgrobacteriumを提供することにより、本発明者ら
は、初めて高頻度のダイズ形質転換を一貫して達成する
ことができた。本発明の方法により、いくつかの異なる
Agrobacterium株でこれらの頻度を得、そしてこの方法
は、Agrobacterium感染に対するダイズ変種の本来の感
受性により最低限の影響を受けるのみである。
形質転換後、外植片は、液体逆選択培地で培養され、
次いて固形の選択培地に移される。このプロセスは、gu
sのようなトランスジェニックマーカーによる同定に関
して、当該分野で記載されるように繰り返される。McCa
beら、（1988）Bio/Technology 6,922−926。シュート
は、トランスジェニック外植片で公知の方法により誘導
される。Wrightら，（1986）Plant Cell Reports 5,150
−154;Barwaleら，（1986）Planta 167,473−481。シュ
ートを切り取り、そしてピログルタミン酸をホルモン富
化成長培地に加えることにより、根が容易に誘導され
る。これは、根はダイズ組織からまれにしか誘導され
ず、従ってホルモンフリー培地でのみ誘導されるという
報告に反する。完全な成熟再生植物は、土壌での温室栽
培に移された後に生成される。
発明の詳細な説明 本発明者らは、gus遺伝子を有するAgrobacterium tu
mefaciensとの共存培養後、ダイズ下胚軸細胞においてE
scherichia coliからβ−グルクロニダーゼ（gus）遺
伝子を発現させた。キメラのgusおよびネオマイシンホ
スホトランスフェラーゼII（npt II）遺伝子を有するバ
イナリープラスミドを中に含んでいる無害化したAgroba
cterium株が、DNA転移のためのベクターであった。共存
培養後、表１に挙げた市販栽培品種由来のダイズ下胚軸
外植片を、100μm/ml硫酸カナマイシンを含有するカル
ス誘導培地（Hincheeら，（1988）Bio/Technology 6,91
5−922）で培養した。形質転換は、下胚軸外植片がgus
活性に対して組織化学的にアッセイされたときに、共存
培養後３週間に評価づけされた。形質転換は、解剖顕微
鏡（倍率10×）を用いて可視化された。これらの形質転
換は、染色植物細胞のセクター（sector）として現れ
た。gus遺伝子のダイズ染色体への安定した組み込み
は、染色セクター内の子孫の細胞集団における遺伝子活
性の保持により示唆された。複数の別個の事象がいくつ
かの外植片で検出され得た。これらは、gus遺伝子を発
現しない細胞により至る所で飛び出した。移植片の組は
共に接種され、共存培養され、選択され、そして組織化
学的にアッセイされた。各組は独立した実験単位であ
り、遺伝的差異（種子ロット内の）、外植片の配置（幼
植物内の）、および外植片の大きさによる変化を補うの
に十分な外植片数である。処理は、その中の外植片で検
出された事象の総数を記録することにより評価された。
この方法を用いて、持続した***が可能な形質転換細胞
のみが評価された。この測定値は、バックグラウンドgu
s活性およびあらゆる残留細菌混入とも明らかに区別さ
れた。
シグナル分子。表１は、８つの異なるダイズ変種の下胚
軸外植片を、アセトシリンゴン（100μＭ）の存在下お
よび不在下で、接種し（30分）そして共存培養した（３
日）実験の結果を示す。接種物は、最終濃度３×108生
存細胞/mlで、アセトシリンゴンを添加してまたは添加
しないで、10mM MESでpH 5.5に緩衝化した液体植物細胞
培地中に、対数期Agrobacteriumを再懸濁することによ
り調製された。外植片は、接種物中で調製され、そこに
30分間保持され、次いで３日間共存培養のために寒天固
形化培地に移された。共存培養は22℃で行った。共存培
養後、外植片は洗浄され、形質転換植物細胞の選択およ
びAgrobacteriaの逆選択のために抗生物質（カナマイシ
ン）を含有する固形培地に移された。継代培養は28℃で
あった。形質転換セクターは、アセトシリンゴンが接種
培地および共存培養培地に添加されたときにのみ生成さ
れた。形質転換セクターは、すべての変種で生成され
た。各変種（48外植片）で生成された形質転換セクター
の平均数は、107であった。形質転換セクター数は、最
低44（Cartter変種）から最高153（9391変種）までの範
囲であった。継代実験では、少数の形質転換セクター
が、アセトシリンゴンを経験しなかった外植片で生成さ
れたが、このような事象の頻度は低すぎて実際に測定す
ることはできなかった。
一晩細菌培養および植物培地を用いる細菌プレ培養期
にアセトシリンゴンを含めても、形質転換頻度は増大し
なかった。単糖類グルコースが接種培地および共存培養
培地に取り入れられたとき、同様の結果が得られた（デ
ータを示さず）。すべての継代実験は、接種培地および
共存培養培地に100μＭでアセトシリンゴンを含んで行
った。
温度。表２は、共通ペトリ皿に接種される下胚軸外植片
を無作為に２組に分けた実験の結果を示す。この組（各
24外植片）を、固形培地上で異なる温度（22℃および28
℃）で、３日間共存培養した。共存培養後、すべての外
植片を28℃で培養した。12回の別個の接種は、9341変種
を用いて、３×108生存細胞/mlのAgrobacteriumで30分
間行われた。22℃で共存培養した24個の外植片で生成さ
れた形質転換セクターの平均数は109であった。28℃形
質転換セクターの平均数は1.4であった。この実験にお
いて、低温での共存培養から生じる形質転換頻度の増大
は、約80倍である。すべての継代共存培養は22℃で行っ
た。
pH。表３は、外植片を異なるpHレベル（5.5、5.75、お
よび6.0）で接種し共存培養した実験の結果を示す。14
回の別個の接種は、各pHで行われた。培地を10μM MES
で緩衝化してpHの安定を確実にした。次いで共存培養
が、同様の緩衝化した固形化培地で３日間行われた。pH
6.0で24外植片で生成された形質転換セクターの平均数
は、pH 5.5（286）またはpH 5.75（288）のいずれかで
生成されたよりも有意に低かった（203）。これは、公
表された結果では、アセトシリンゴンを用いたときに、
Glycine maxにおいてpHの表示効果が全く示されなかっ
たことに反する。Godwinら，（1991）Plant Cell Rep.
9,671−675。すべての継代接種および共存培養は、pH
5.5に調整され、10μM MESで緩衝化された培地で行っ
た。
接種物における生存細菌濃度。表４は、外植片に異なる
濃度の細菌を接種した２つの実験の結果を示す。これら
の実験に関し、対数期の一晩細菌培養物を沈降し、次い
で適切な濃度への順次希釈により再懸濁した。各実験に
おいて、３つの別個の細菌希釈系列を共通一晩培養から
調製した。前述の実験におけるように、48個の外植片を
各接種物において調製し、そこで30分間保持し、固形培
地で３日間共存培養した。48個の外植片の各接種に対し
て生成された形質転換セクターの総数を記録した。最初
の３つの接種濃度:3×107、108、および３×108細胞/ml
について、より多くの形質転換セクターが、より多い細
菌により生成された。これは、両実験およびすべての希
釈系列に関していえる。
より高い接種濃度（109細胞/ml）では、形質転換頻度
の増大は検出されなかった。この濃度で生成されたセク
ターの平均数は、それ以下の濃度の平均数と有意に違わ
なかった。従って、投与量／反応プロフィールは、細菌
数が制限される濃度から細菌が飽和である濃度まで、３
×108細胞/mlで変化する。
所定の濃度で生成される形質転換体の絶対数は、実験
間でかなり変化する。各実験においてプラトーが見かけ
上達成されるので、Agrobacteriaの数よりもむしろその
状態が限定要因となるようである。異なるダイズ外植片
調製物が形質転換の能力により変化したことも考えられ
る。第三に考えられることは、接種の真の長さが実験間
で変化していることである。これはありそうにないこと
であるが、実験は、所定の濃度を飽和させるのに十分な
接種期間を規定して行った。30分の期間は、最大数の事
象を生じるように決定され、さらに時間を加えても何の
効果も生じなかった（データ示さず）。
連続接種。Agrobacteriaをダイズ細胞に感染させる試み
において、48個のダイズ下胚軸外植片（9341変種）の組
を、５×108生存細胞/mlで細菌を30分間、１回または２
回接種した実験を行った。すべての外植片は３日間共存
培養した。結果は表５に示す。形質転換セクターの平均
数は、２回処理（315）の方が１回処理（286）よりも有
意に高かった。連続接種により、さらに形質転換は生じ
得ないようである。形質転換可能なダイズ細胞プールの
真の飽和は１回接種により達成され、そして生存細菌数
は、接種が非常に高濃度でなされたときであっても制限
されたままである。しかし、形質転換に至る連合の大部
分は、最初の30分接種で生じている。
感受性および耐性ダイズ遺伝子型。表１に示した結果
は、試験したすべてのダイズ変種は、本発明の方法を用
いてAgrobacteriumにより形質転換され得ることを示し
た。Agrobacterium媒介形質転換に対するダイズ変種の
相対的感受性を決定するために、推定耐性変種（Corsoy
79およびCentury）の交雑組み合わせ検定を、感受性パ
ートナーとしてPeking変種を用いて行った。接種は、異
なる変種の外植片が共通の接種物において調製され得る
ように、ステンレスメッシュ分離器（0.5mm孔）を有す
るペトリ皿で行った。「耐性」変種の24個の外植片を、
等しい数の「感受性」外植片を含む各12個の別個のペト
リ皿中で調製した。結果は表６に示す。変種Pekingは、
変種Centuryの約２倍の頻度および変種Corsoy 79の約３
倍の頻度で形質転換された。変種に依存する感受性がこ
れらの実験により確認されているが、その程度（２〜３
倍）は小さいので、多様なダイズ生殖質の形質転換にAg
robacteriumを用いることに関して重大な示唆を与える
ほどではない。
Agrobacterium株。２つの互いの無害化されたAgrobacte
rium株と、オクトパイン（octopine）株であるLBA4404
とを、ダイズ下胚軸細胞の形質転換を媒介する能力につ
いて比較した；それらは、L,L−スクシナモパイン（suc
cinamopine）株EHA 101およびノパリン（nopaline）株C
58−pz707であった。表７は、これらの各株の濃度を増
大させて、Williams 82の下胚軸外植片への接種に用い
る実験の結果を示す。48個の外植片を各濃度の各株に接
種した。LBA4404をコントロールとして３×108生存細胞
/mlで含有させた。両方の株で形質転換セクターが生じ
た。両方の場合において、形質転換頻度は、接種物にお
ける細菌濃度に依存し、細菌数が多いほど形質転換は多
く生じた。試験したレベルでは、形質転換において、ど
ちらもLBA4404と比較して相当な変化はみられなかっ
た。
以下の実施例は、本発明に従って種々の適用を説明す
るが、決して本発明の範囲を限定する意図はない。
実施例１ ダイズ（Glycine max）種子（Pioneeer変種9341）
は、鐘状ガラス容器中で放出された塩素ガスに曝すこと
により表面を滅菌した。ガスを、3.5mlの塩酸（34〜37
％w/w）を100mlの次亜塩素酸ナトリウム（5.25％w/w）
に加えることにより作製した。約１立方フィートの容量
のコンテナ中で16〜20時間曝した。表面滅菌した種子
を、室温でペトリ皿中に保存した。種子は、植物成長調
節剤を含まないGamborgに従う1/10強度の寒天固形化培
地［最小有機物を含むB5基本培地、Sigma Chemical C
o.,カタログ番号G5893、0.32gm/L;ショ糖、0.2％w/vお
よび２−［Ｎ−モルホリノ］エタンスルホン酸（ME
S）、3.0mM］のプレートに置き、28℃で昼の長さを16時
間にして、約20μEm2S1の弱い白色蛍光照明をあてて培
養することにより、発芽させた。３、４日後、主を共存
培養用に調製し得た。種子の外皮を取り除き、伸長して
いる幼根を子葉の下部３〜4mmで取り除いた。いくつか
のペトリ皿のそれぞれにつき10個の得られた種子を保持
した。
実施例２ 1.0μg/mlのテトラサイクリンを含む最小Ａ培地で対
数期へと増殖した、バイリープラスミドp12GUSBN17（DP
1816）またはp12−4X（DP1813）を有するAgrobacterium
tumefaciens LBA 4404株の終夜培養物をプールし、55
0nmでの吸光度測定を行った。かなりの容量の培養物
を、沈澱して1.0×1010細胞と2.0×1010細胞との間の細
胞が各管に集められるように、15mlの円錐形遠沈管に入
れた。ここでO.D.550 1.0＝1.4×109細胞/mlである。沈
澱は6000gで10分間の遠心により得た。遠心後、上清を
デカントし、接種物が必要になるまで（しかし１時間は
超えない）管を室温で保持した。
実施例３ 接種を、各プレートの種子を新たに再懸濁したAgroba
cteriumのペレットで処理するようなバッチで行った。
ペレットを20mlの接種培地中に一度に再懸濁した。接種
培地は、B5塩（G5893）、3.2gm/L;ショ糖、2.0％w/v;6
−ベンジルアミノプリン（BAP）、44μM;インドール酪
酸（IBA）、0.5μM;アセトシリンゴン（AS）、100μＭ
からなり、10mMのMESでpH5.5に緩衝化した。再懸濁はボ
ルテックスにより行った。次いで接種物を、調製した種
子を含むペトリ皿に注ぎ、子葉節を外科用メスで浸軟さ
せた。これは、２つの子葉全体を保護する茎頂（shoot
apex）を通り、経線部分で半分に種子を分けることによ
り行った。次いで２つの半分の茎頂は、外科用メスでそ
れらを取り除くことによりそれぞれの子葉を分解した。
次いで子葉節を、外科用メスで対称な尖にそって切れ目
を繰り返し付けることにより浸軟した。外植片から軸外
側をまったく切断しないように気を付ける。20個の外植
片をざっと５分間で調製し、次いで室温で30分間撹拌せ
ずにインキュベートした。追加のプレートをこの間に調
製した。外植片を0.2％w/vのGelrite（Merck ＆ Co.,In
c.）で固形化させた同じ培地のプレートに移して30分後
に、この外植片を軸方向側を上にして培地の表面層に埋
め、22℃で３日間、約20μEm2S1の弱い白色蛍光下で培
養した。
実施例４ ３日後に、外植片を液体逆選択培地に移した。逆選択
培地は、B5塩（G5893）、3.2gm/L;ショ糖、2.0％w/v;BA
P、5.0μM;IBA、0.5μM;バンコマイシン、200μg/ml;セ
フォタキシム、500μg/mlからなり、3mMのMESでpH5.7に
緩衝化した。10個の外植片を各ペトリ皿中で、一定のゆ
っくりした旋回撹拌しながら室温で４日間洗浄した。逆
選択培地は４回交換した。
実施例５ 次いで外植片を寒天固形化選択培地に取り出した。選
択培地は、B5塩（G5893）、3.2gm/L;ショ糖、2.0％w/v;
BAP、5.0μM;IBA、0.5μM;硫酸カナマイシン、50μg/m
l;バンコマイシン、100μg/ml;セフォタキシム、30μg/
ml;チメンチン（timentin）、30μg/mlからなり、3.0mM
のMESでpH5.7に緩衝化した。選択培地を0.3％w/vのSeaK
emアガロースで固形化した。外植片を軸方向を下にして
培地に埋め、28℃で昼の長さを16時間にして60〜80μEm
2S1の弱い白色蛍光照明をあてて培養した。
実施例６ ２週間後、外植片を再び旋回シェーカー上で液体培地
で洗浄した。今回は、洗浄を50μg/mlの硫酸カナマイシ
ンを含む逆選択培地中で終夜で行った。次の日、外植片
を寒天固形化選択培地に取り出した。再びそれらを培地
中に軸方向を下に埋めた。培養をもう２週間、前のよう
に行った。
実施例７ １カ月後選択培地上で、経質転換組織は、バックグラ
ウンドの退色したあまり健常ではない組織に対して緑色
セクターの再生組織として認められるようになった。緑
色セクターのない外植片を放棄し、緑色セクターのある
外植片を伸長培地（elongation medium）に移した。伸
長培地は、B5塩（G5893）、3.2gm/L;ショ糖、2.0％w/v;
IBA、3.3μM;ジベレリン酸、1.7μM;バンコマイシン、1
00μg/ml;セフォタキシン、30μg/ml;およびチメンチ
ン、30μg/mlからなり、3.0mMのMESでpH5.7に緩衝化し
た。伸長培地は、0.2％w/vのgelriteで固形化した。そ
れらは、軸方向を上にして埋め、前のように培養した。
培養を、この培地で２週間おきに新鮮なプレートに移し
て継続した。シュートが0.5cmの長さになった場合、そ
れを根元で切除し、13×100mmの試験管中の発根培地に
置いた。発根培地は、B5塩（G5893）、3.2gm/L;ショ
糖、15gm/L;ニコチン酸、20μM;ピログルタミン酸（PG
A）、900mg/L、およびIBA、10μＭからなっていた。そ
れは、3.0mMのMESでpH5.7に緩衝化し、0.2％w/vのGelri
teで固形化した。10日後、シュートをIBAまたはPGAを含
まない同培地に移した。シュートを発根させ、前と同じ
環境条件下でこれらの管内に保持した。
実施例８ 根系がうまく樹立した場合、幼植物をプラントコン
（plant con）（ICN Biomedicals,Inc.,カタログ番号26
−720＆１−02）の中の滅菌した混合土に移した。温
度、日長、および光強度は前のまま維持した。これらの
条件下で、再生植物は、育ちが良く（小さいが）ほとん
ど正常の植物となった。それらの根系が再びうまく樹立
された場合、プラントコンのコーナーを切り落とし、植
物を環境チャンバーまたは温室で徐々に寒気に慣らし
た。最終的に、それらは混合土に鉢上げし、温室で成熟
して種子を生じるまで生育させた。
───────────────────────────────────────────────────── フロントページの続き (56)参考文献 Ｐｌａｎｔ Ｃｅｌｌ Ｒｅｐｏｒｔ ｓ ９（1991）Ｐ．671−675 (58)調査した分野(Int.Cl.6，ＤＢ名) C12N 15/09 C12N 5/00 C12N 1/21 A01H 1/00 ＢＩＯＳＩＳ（ＤＩＡＬＯＧ) ＷＰＩＬ（ＤＩＡＬＯＧ)

Claims (7)

    (57)【特許請求の範囲】
  1. 【請求項１】以下の工程を包含する、遺伝子型に依存し
    ない、トランスジェニックダイズ植物の生成方法： a. 発芽させたダイズ種子の下胚軸または培養子葉節由
    来の外植片を、108から３×108細胞/mlの濃度のキメラ
    遺伝子を含むAgrobacterium種細胞と共に、アセトシリ
    ンゴン、α−ヒドロキシアセトシリンゴン、アセトバニ
    ロン、シリンガアルデヒド、シリンガ酸、シナピン酸、
    およびそれらの混合物からなる群から選択されるシグナ
    ル化合物の存在下で共存培養する工程； b. 18〜28℃の共存培養温度を維持する工程；および c. 植物培養培地pHをpH6.0未満にまで低下させること
    により該Agrobacteriumの毒性を誘導する工程。
  2. 【請求項２】前記共存培養がAgrobacteriumの順次接種
    により行われる、請求項１に記載の方法。
  3. 【請求項３】前記キメラ遺伝子が種子貯蔵タンパク質遺
    伝子である、請求項１に記載の方法。
  4. 【請求項４】前記キメラ遺伝子がBertholletia excelsa
    由来の2S貯蔵タンパク質である、請求項１に記載の方
    法。
  5. 【請求項５】子葉節からダイズ植物を再生する工程をさ
    らに包含する、請求項１から４に記載の方法であって、
    以下の工程を包含する、方法： a. 該節を分割する工程； b. 該分割した節を、栄養培地上でカルス組織が発達す
    るまで培養する工程であって、該組織がシュートを含
    む、工程；および c. 該カルスから該シュートを切り取り、そしてホルモ
    ンおよびピログルタミン酸を含有する栄養培地上で該シ
    ュートを発根させて幼植物を形成する工程。
  6. 【請求項６】前記発根培地がGamborg培地である、請求
    項５に記載の方法。
  7. 【請求項７】前記幼植物が、土を含む培地に移され、該
    幼植物から完全植物が生じるまで該培地上で生育され
    る、請求項５に記載の方法。
JP6504746A 1992-07-27 1993-07-26 培養ダイズ細胞のａｇｒｏｂａｃｔｅｒｉｕｍ媒介形質転換の改良法 Expired - Lifetime JP2952041B2 (ja)

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