JP5721209B2 - 根こぶ病抵抗性アブラナ科植物の作出方法 - Google Patents
根こぶ病抵抗性アブラナ科植物の作出方法 Download PDFInfo
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Classifications
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- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
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- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
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- 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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Description
効率的な育種を推進するためには、遺伝子組換え技術を用いて抵抗性遺伝子の導入によって育成を図る方法と、交配が容易な種には、遺伝子座周辺のゲノム情報を活用してマーカー選抜により抵抗性品種を育成する方法がある。本発明者らは、根こぶ病抵抗性遺伝子を単離し、遺伝子組み換えおよびマーカー選抜により効率的な根こぶ病抵抗性のアブラナ科植物を作出する方法を提供する。
〔1〕 以下の(a)から(d)のいずれかに記載の、根こぶ病菌抵抗性を有するポリヌクレオチド;
(a)配列番号:2に記載のアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(b)配列番号:1に記載の塩基配列のコード領域を含むポリヌクレオチド、
(c)配列番号:2に記載のアミノ酸配列において1または複数のアミノ酸が置換、欠失、付加、および/または挿入されたアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(d)配列番号:1記載の塩基配列の相補鎖にストリンジェントな条件でハイブリダイズするポリヌクレオチド。
〔2〕 植物細胞で発現可能なプロモーター領域の下流に、〔1〕に記載のポリヌクレオチドが機能的に結合したベクター。
〔3〕 〔2〕に記載のベクターが導入された形質転換植物細胞。
〔4〕 〔3〕に記載の形質転換植物細胞から再生された、根こぶ病菌抵抗性活性を有する植物体。
〔5〕 〔4〕に記載の植物体の子孫またはクローンである、根こぶ病菌抵抗性活性を有する植物体。
〔6〕 〔4〕または〔5〕に記載の根こぶ病菌抵抗性活性を有する植物体の繁殖材料。
〔7〕 配列番号:1もしくは配列番号:3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を検出する工程を含む、被検植物体または被検繁殖媒体の根こぶ病菌抵抗性を判定する方法。
〔8〕 以下の(i)から(iii)の工程を含む、〔7〕に記載の判定方法;
(i)被検植物体または被検繁殖媒体からDNA試料を調製する工程、
(ii)該DNA試料から配列番号:1もしくは配列番号:3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅する工程、
(iii)根こぶ病菌抵抗性の植物体または繁殖媒体から、配列番号:1もしくは3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅したDNA断片と、工程(ii)において増幅したDNA断片の分子量または塩基配列を比較する工程。
〔9〕 〔7〕または〔8〕に記載の判定方法により、根こぶ病菌抵抗性遺伝子を有する植物体またはその種子を選抜する方法。
〔10〕 以下の(i)および(ii)の工程を含む、根こぶ病菌抵抗性活性を有する植物体またはその種子の製造方法;
(i)〔2〕に記載のベクターを植物細胞に導入する工程、および
(ii)前記工程(i)においてベクターが導入された形質転換植物細胞から植物体を再生する工程。
〔11〕 〔1〕に記載のポリヌクレオチドを植物細胞内で発現させる工程を含む、植物体もしくはその種子に根こぶ病抵抗性活性を付与する方法。
〔12〕 〔1〕に記載のポリヌクレオチド、または〔2〕に記載のベクターを植物細胞へ導入する工程を含む、〔11〕に記載の方法。
〔13〕 以下の(a)および(b)の工程を含む、〔11〕に記載の方法;
(a)〔1〕に記載のポリヌクレオチドを有する植物と他の植物とを交雑させる工程、および
(b)前記ポリヌクレオチドを有する植物体を選抜する工程。
〔14〕 植物がアブラナ科植物である、〔7〕〜〔13〕のいずれかに記載の方法。
〔15〕 〔7〕〜〔14〕のいずれかに記載の方法によって取得される植物体またはその種子。
〔16〕 人為的に作製された植物体またはその種子であって、〔1〕に記載のポリヌクレオチドを有し根こぶ病菌抵抗性活性を有することを特徴とする植物体またはその種子。
〔17〕 植物がアブラナ科植物である、〔15〕もしくは〔16〕に記載の植物体またはその種子。
〔18〕 配列番号:1または配列番号:3に記載の塩基配列とストリンジェントな条件で特異的にハイブリダイズし、少なくとも15ヌクレオチドの鎖長を有するオリゴヌクレオチドを含む、被検植物の根こぶ病菌抵抗性活性を検出するためのプライマー。
〔19〕 配列番号:1または配列番号:3に記載の塩基配列とストリンジェントな条件で特異的にハイブリダイズし、少なくとも15ヌクレオチドの鎖長を有するオリゴヌクレオチドを含む、被検植物の根こぶ病菌抵抗性活性を検出するためのプローブ。
(a)配列番号:2に記載のアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(b)配列番号:1に記載の塩基配列のコード領域を含むポリヌクレオチド、
(c)配列番号:2に記載のアミノ酸配列において1または複数のアミノ酸が置換、欠失、付加、および/または挿入されたアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(d)配列番号:1および配列番号:3記載の塩基配列の相補鎖にストリンジェントな条件でハイブリダイズするポリヌクレオチド。
本発明のベクターとしては、組換えタンパク質の生産に用いる上記ベクターの他、形質転換植物体作製のために植物細胞内で本発明のポリヌクレオチドを発現させるためのベクターも含まれる。本発明の好ましい態様として例えば、植物細胞で発現可能なプロモーター領域の下流に、本発明のポリヌクレオチドが機能的に結合したベクターを挙げることができる。例えば、植物細胞で転写可能なプロモーター配列と転写産物の安定化に必要なポリアデニレーション部位を含むターミネーター配列を含むことができる。植物細胞の形質転換に用いられるベクターとしては、該細胞内で挿入遺伝子を発現させることが可能なものであれば特に制限はない。例えば、植物細胞内での恒常的な遺伝子発現を行うためのプロモーターを有するベクターや外的な刺激により誘導的に活性化されるプロモーターを有するベクターを用いることも可能である。
(i)被検植物体または被検繁殖媒体からDNA試料を調製する工程、
(ii)該DNA試料から配列番号:1もしくは配列番号:3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅する工程、
(iii)根こぶ病菌抵抗性の植物体または繁殖媒体から、配列番号:1もしくは3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅したDNA断片と、工程(ii)において増幅したDNA断片の分子量または塩基配列を比較する工程。
(i) 本発明のベクターを植物細胞に導入する工程、および
(ii) 前記工程(i)においてベクターが導入された形質転換細胞から植物体を再生する工程。
(a)本発明のポリヌクレオチドを有する植物と他の植物とを交雑させる工程、および
(b)前記ポリヌクレオチドを有する植物体を選抜する工程。
(a)配列番号:1および配列番号:3に記載の塩基配列の全配列もしくはその一部の配列を増幅するためのオリゴヌクレオチドプライマー
(b)配列番号:1および配列番号:3に記載の塩基配列とストリンジェントな条件でハイブリダイズし、少なくとも15ヌクレオチドの鎖長を有するオリゴヌクレオチドプローブ
根こぶ病抵抗性遺伝子をマップベースクローニングにより単離するためにヨーロッパ飼料用カブ「Siloga」由来の抵抗性系統G004のBACライブラリーを構築した。このライブラリーの平均インサート長は67.4 kb、そのサイズは約38,400クローンで、550 MBと推定されるB.rapa のゲノム(Arumuganathan K, Earle ED., Plant Mol Biol Rep, (1991) 9 (3): 208-219.)の4.7倍に相当した。Crr1を有するBACクローンを単離するため、連鎖するマーカーの中からCrr1に最も連鎖するマーカーであるBSA7(図1、Suwabe, K. et al., Genetics, (2006) 173: 309-319)を起点に染色体歩行を行った。
BSA7を起点としてCrr1周辺領域がBACクローンでカバーされ、これらのBACの末端配列を罹病性品種PL7とG004間で比較することにより多型を示す複数のマーカーが得られた。BACクローンの末端配列から得られたマーカー遺伝子型と抵抗性遺伝子の有無の比較により、B355H7とB359C3の末端配列から作成したマーカーがCrr1に最も近傍なマーカーであった。
F2 No.1075 rr Rr
F2 No.764 Rr rr
F2 No.572 RR Rr
B355H7とB359C3の間約8 kbのDNA配列を決定するためB355H7のショットガンクローンを作製した。抽出したプラスミド断片のDNA配列をT7とReverse Primerを用いて定法に従い決定し、DNAシーケンスアセンブルソフトウェアSEQUENCHERver.2(日立ソフトウェアエンジアリング、東京)を用いて1つのシーケンスとした。この領域の中でタンパク質をコードするオープンリーディングフレーム(ORF)を遺伝情報処理ソフトウェア GENETYX(ゼネティックス、東京)を用いて検索した。決定したG004 配列情報を基に適当な位置にプライマーを設計し、罹病性のPL7を鋳型にしてDNA断片を増幅し、PL7の当該領域の塩基配列を決定した。PL7とG004 との間でDNA配列比較を行い、挿入欠失配列や一塩基多型の有無を調べた。結果、抵抗性系統G004のB355H7とB359C3間のDNA断片長は7,995 bpであり、この配列の中には4個のORFが推定された(図4、配列番号:3)。
(1)植物細胞導入用ベクターの構築
・レタスユビキチン遺伝子をプロモーターに用いたベクターの構築
レタスからクローニングしたユビキチン遺伝子プロモーターとターミネーターの間にGUS遺伝子が挿入されたプラスミドpUC198UGU(福岡浩之博士より分譲)をSmaIとEcoICRIで切断し、予めSmaIで処理したCrr1 cDNA配列を挿入してpUC198UCrr1Uを構築した(図9)。レタスユビキチンプロモーター、Crr1 cDNA、レタスユビキチンターミネーターのカセットをAscIで切り出し、pPZP202(Hajdukiewicz P. et al., Plant Mol Biol, (1994) 25: 989-994)より改変したバイナリベクターpZK3B(農研機構 中央農業研究センター・黒田昌治博士より分譲)に挿入しpUbp-Crr1_ZK3Bを構築した(図9)。
シロイヌナズナでの根こぶ病検定は、Jubault M. et al., Theor.Appl.Genet.,(2008) 117:191-202の方法を改変して行った。カナマイシン耐性の植物体を1ポットあたり9個体になるように園芸倍土(TM-1, タキイ種苗)に移植し、3日後に休眠胞子液(1.0 x 106胞子/ml)2 mlを植物の株元に灌注した。22℃、14時間日長条件で栽培し、接種3週間後に根の病徴程度を調査した。発病程度は、0(無病徴)、1(側根に小さなコブ)、2(側根に少し大きなコブ)、3(主根に大きなコブ、胚軸も肥大)の4段階で評価した。
BRMS-173とBRMS-088を用いたマーカー選抜では、マーカー座とCrr1座との間で組換えを起こす個体がわずかな確率ではあるが生じる。これを回避するためには抵抗性遺伝子そのものをマーカーに用いることが想定される。配列番号:3にあるとおり、抵抗性と罹病性系統間でCrr1のゲノム配列を比較すると挿入・欠失配列が多く存在する。これらを用いて抵抗性個体を選抜するマーカーとしての有用性を検討した。
※1:Crr1とCrr2とは別々の染色体に座乗しており、B359C3はCrr1遺伝子、BRMS-096はCrr2に連鎖するDNAマーカーである。 rr:罹病性親、Rr:ヘテロ型、RR:抵抗性親型
※2:発病指数;0:無病徴、1:側根に小さなコブ、2:側根に連続したコブ、3:主根にコブ
※3:平均発病指数: Σ(発病指数*個体数)/(総個体数)
Claims (16)
- 以下の(a)から(d)のいずれかに記載の、根こぶ病菌抵抗性を有するポリヌクレオチド;
(a)配列番号:2に記載のアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(b)配列番号:1に記載の塩基配列のコード領域を含むポリヌクレオチド、
(c)配列番号:2に記載のアミノ酸配列において1または複数のアミノ酸が置換、欠失、付加、および/または挿入されたアミノ酸配列からなるタンパク質をコードするポリヌクレオチド、
(d)配列番号:1記載の塩基配列の相補鎖に0.2×SSC、0.1%SDS、65℃の条件でハイブリダイズするポリヌクレオチド。 - 植物細胞で発現可能なプロモーター領域の下流に、請求項1に記載のポリヌクレオチドが機能的に結合したベクター。
- 請求項2に記載のベクターが導入された形質転換植物細胞。
- 請求項3に記載の形質転換植物細胞から再生された、根こぶ病菌抵抗性活性を有する植物体。
- 請求項4に記載の植物体の子孫またはクローンである、根こぶ病菌抵抗性活性を有する植物体。
- 請求項4または5に記載の根こぶ病菌抵抗性活性を有する植物体の繁殖材料。
- 配列番号:1もしくは配列番号:3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を検出する工程を含む、被検植物体または被検繁殖媒体の根こぶ病菌抵抗性を判定する方法。
- 以下の(i)から(iii)の工程を含む、請求項7に記載の判定方法;
(i)被検植物体または被検繁殖媒体からDNA試料を調製する工程、
(ii)該DNA試料から配列番号:1もしくは配列番号:3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅する工程、
(iii)根こぶ病菌抵抗性の植物体または繁殖媒体から、配列番号:1もしくは3に記載の塩基配列、または該塩基配列の部分配列もしくは周辺配列を含むDNA領域を増幅したDNA断片と、工程(ii)において増幅したDNA断片の分子量または塩基配列を比較する工程。 - 請求項7または8に記載の判定方法により、根こぶ病菌抵抗性遺伝子を有する植物体またはその種子を選抜する方法。
- 以下の(i)および(ii)の工程を含む、根こぶ病菌抵抗性活性を有する植物体またはその種子の製造方法;
(i)請求項2に記載のベクターを植物細胞に導入する工程、および
(ii)前記工程(i)においてベクターが導入された形質転換植物細胞から植物体を再生する工程。 - 請求項1に記載のポリヌクレオチドを植物細胞内で発現させる工程を含む、植物体もしくはその種子に根こぶ病抵抗性活性を付与する方法。
- 請求項1に記載のポリヌクレオチド、または請求項2に記載のベクターを植物細胞へ導入する工程を含む、請求項11に記載の方法。
- 以下の(a)および(b)の工程を含む、請求項11に記載の方法;
(a)請求項1に記載のポリヌクレオチドを有する植物と他の植物とを交雑させる工程、および
(b)前記ポリヌクレオチドを有する植物体を選抜する工程。 - 植物がアブラナ科植物である、請求項7〜13のいずれかに記載の方法。
- 配列番号:1または塩基配列:3に記載の塩基配列とストリンジェントな条件で特異的にハイブリダイズし、少なくとも15ヌクレオチドの鎖長を有するオリゴヌクレオチドを含む、被検植物の根こぶ病菌抵抗性活性を検出するためのプライマー。
- 配列番号:1または塩基配列:3に記載の塩基配列とストリンジェントな条件で特異的にハイブリダイズし、少なくとも15ヌクレオチドの鎖長を有するオリゴヌクレオチドを含む、被検植物の根こぶ病菌抵抗性活性を検出するためのプローブ。
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JP4366494B2 (ja) * | 2002-10-16 | 2009-11-18 | 独立行政法人農業・食品産業技術総合研究機構 | アブラナ科植物根こぶ病に対する抵抗性遺伝子検出用マイクロサテライトマーカー、およびその利用 |
JP4469992B2 (ja) | 2003-12-16 | 2010-06-02 | 三重県 | 根こぶ病抵抗性ナタネ類の選抜マーカー |
JP5164150B2 (ja) | 2008-01-30 | 2013-03-13 | 独立行政法人農業生物資源研究所 | アブラナ科野菜根こぶ病菌による根こぶ形成時特異的プロモーター |
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2010
- 2010-09-22 JP JP2010211689A patent/JP5721209B2/ja active Active
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2011
- 2011-09-22 CA CA2811878A patent/CA2811878A1/en not_active Abandoned
- 2011-09-22 WO PCT/JP2011/071554 patent/WO2012039445A1/ja active Application Filing
- 2011-09-22 EP EP11826890.3A patent/EP2620499A4/en not_active Withdrawn
- 2011-09-22 US US13/825,783 patent/US20130254929A1/en not_active Abandoned
Also Published As
Publication number | Publication date |
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WO2012039445A1 (ja) | 2012-03-29 |
CA2811878A1 (en) | 2012-03-29 |
EP2620499A4 (en) | 2014-04-16 |
JP2012065567A (ja) | 2012-04-05 |
US20130254929A1 (en) | 2013-09-26 |
EP2620499A1 (en) | 2013-07-31 |
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