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{"target":"https://pubannotation.org/docs/sourcedb/FA@AikoHIRAKI/sourceid/16954_","sourcedb":"FA@AikoHIRAKI","sourceid":"16954_","text":"〈タイトル〉根から長距離を輸送されたサイトカイニンとその前駆体は地上部の成長の制御において異なる役割を担う\r\n\r\n〈著 者〉大薄麻未・榊原 均\r\n〈著者所属〉名古屋大学大学院生命農学研究科 生物機構・機能科学専攻生物化学研究分野\r\n〈著者email〉asami.yoshizumi@utoronto.ca(大薄麻未),sakaki@agr.nagoya-u.ac.jp(榊原 均)\r\n\r\n〈対象論文〉\r\nSystemic transport of trans-zeatin and its precursor have differing roles in Arabidopsis shoots.\r\nAsami Osugi, Mikiko Kojima, Yumiko Takebayashi, Nanae Ueda, Takatoshi Kiba, Hitoshi Sakakibara\r\nNature Plants, 3, 17112 (2017)\r\n\r\n〈要 約〉\r\n 植物の個体は環境に最適化された形態をとるため,器官のあいだで情報をやりとりしておのおのの器官の成長のバランスを制御する.植物ホルモンのサイトカイニンは器官のあいだのシグナル分子のひとつである.サイトカイニンの前駆体であるトランスゼアチンリボシドは道管を輸送されるサイトカイニン関連化合物の主要な成分であり,根の窒素栄養の環境に応答してその濃度が変動することから,根の栄養環境を地上の器官に伝達するシグナル分子であると考えられてきた.一方,採取した道管液からは活性型のサイトカイニンであるトランスゼアチンが微量に検出されるが,その輸送および作用についてはほとんど注目されていなかった.この研究において,道管にはサイトカイニンとしてトランスゼアチンリボシドとトランスゼアチンの2種類の輸送形態の存在が明らかにされた.そして,サイトカイニン生合成系の遺伝子や輸送系の遺伝子の変異体を組み合わせた接木植物を詳細に解析することにより,根から長距離を輸送されたトランスゼアチンはおもに葉の面積の制御にかかわるのに対し,トランスゼアチンリボシドは葉の形成の速度をも制御することが明らかにされた.\r\n\r\nはじめに\r\n 植物は機能的に分化した複数の器官から構成されており,それぞれ異なる環境の変化にさらされている.たとえば,主要な光合成器官である葉は光や二酸化炭素の濃度の変化にさらされるのに対し,地下器官である根は栄養環境や土壌の水分含量の変動にさらされる.このような局所的な環境の変化をほかの器官に伝達し個体のレベルで調和のとれた形態をとるために,植物はさまざまなシグナル分子を長距離にわたり輸送し器官のあいだでコミュニケーションをとっている.\r\n 水,栄養,光合成産物を全身へ分配するための道管および師管は,器官のあいだのシグナル分子の長距離の輸送も担うことが知られている.道管は根から地上部へのシグナル分子の輸送を担うのに対し,師管はおもに成熟葉(ソース器官)から花芽,根,未熟葉(シンク器官)にむけた輸送を担う.これまでに,RNA,タンパク質,代謝産物,植物ホルモンなど,さまざまな分子が器官のあいだのシグナル分子として報告されている.サイトカイニンは,葉の成長の促進,地上部の幹細胞組織である茎頂分裂組織の維持,花芽の形態形成,根の成長の抑制などの作用をもつ植物ホルモンであるが1,2),器官のあいだのシグナル分子としての役割ももつ.サイトカイニンの前駆体であるトランスゼアチンリボシドは道管液において検出されるサイトカイニン関連化合物の大部分をしめることから,根から地上部へのサイトカイニンの輸送型であることが示唆されている3).トランスゼアチンリボシドはサイトカイニンの前駆体であり,作用するまえにヌクレオチドに変換され,そののち,LOGとよばれる酵素により活性型のサイトカイニンであるトランスゼアチンへと変換される.道管を輸送されるトランスゼアチンリボシドの量は根の窒素栄養の環境に応答して制御されることから,トランスゼアチンリボシドは根の栄養環境を地上部へと伝達するシグナル分子であると考えられてきた.一方,道管液にはトランスゼアチンリボシドのほか微量のトランスゼアチンも検出されるが,このトランスゼアチンは実際に長距離輸送されるのか,また,どのような形質に対し作用するのかについては明らかにされていなかった.\r\n\r\n1.シグナル分子の長距離輸送の研究への接木技術の利用\r\n 近年,植物における長距離輸送される分子の解析において,接木技術が強力なツールとして用いられている.接木は植物のもつ高い脱分化能および再分化能を利用したもので,異なる2つの植物体の切断面を接触させることにより癒合を促進しひとつの個体を生み出すものである.2つの植物体のうち,上部を穂木,下部を台木とよぶ.接木は園芸分野においてよく利用されており,たとえば,土壌に由来する病害に弱い園芸作物の穂木と病害に強い植物種の台木とを接木することにより園芸作物の耐病性を強めることができる.長距離輸送される分子の解析に接木技術を用いる例として,GFPなどのタグを付加した遺伝子をもつ形質転換体と野生型の植物体とを接木し,野生型に由来する器官においてタグを検出することによりRNAやタンパク質の長距離輸送を証明できる(図1a).また,その表現型を観察することにより長距離輸送される分子の機能を検証できる.接木植物を遺伝学に解析することにより内在性の分子の長距離輸送を検証することもできる(図1b).ある遺伝子の欠損変異体においては,これによりコードされるRNAおよびタンパク質が欠失するだけでなく,それにより合成される代謝産物やシグナル分子などの存在量も影響をうける.そこで,変異体と野生型とを接木し,変異体に由来する器官においてこれら代謝産物やシグナル分子の存在量が回復するかどうかを解析することにより,野生型に由来する器官から変異体に由来する器官へと長距離輸送されるかどうかを検証することができる.さらに,変異体に由来する器官における表現型を解析することにより長距離輸送された分子の機能を解析できる.この研究においては,活性型サイトカイニンの長距離輸送を検証するために,また,長距離輸送された活性型サイトカイニンの生理作用を検証するために,シロイヌナズナのサイトカイニン生合成系の変異体を用いた接木植物を作出し解析した.\r\n\r\n2.トランスゼアチンは根から地上部へと輸送され作用する\r\n 最新の解析技術を用いてシロイヌナズナの道管液におけるサイトカイニンおよび関連化合物の組成を解析したところ,これまでの報告どおり,サイトカイニン前駆体であるトランスゼアチンリボシドがもっとも豊富で全体の約80%をしめ,活性型サイトカイニンであるトランスゼアチンが約20%をしめた.そこで,道管液から検出されるトランスゼアチンが根から地上部へと長距離輸送されるかどうか検証するため,安定同位体により標識したトランスゼアチンを用いてトレース実験を行った.標識したトランスゼアチンを野生型のシロイヌナズナの根に投与し,地上部を採取して解析した結果,地上部においても標識されたトランスゼアチンが検出された.しかし,トランスゼアチンは生体において前駆体であるトランスゼアチンリボシドやそのほかの派生体へと容易に変換されることから,投与したトランスゼアチンがそのまま輸送されたとはかぎらない.そこで,接木技術を利用することにした.\r\n LOGはサイトカイニンの活性化を担う酵素であり,LOGをコードする遺伝子の多重変異体であるlog七重変異体においてトランスゼアチンの生合成量はいちじるしく低下する4).このlog七重変異体を穂木,野生型を台木とした接木植物において同様のトレース実験を行ったところ,野生型と同様に,地上部において標識されたトランスゼアチンが検出された.また,根から地上部へのサイトカイニンの輸送を担うことが報告されているABCG14遺伝子5) の変異体であるabcg14変異体を用いて,log七重変異体を穂木,abcg14変異体を台木にした接木植物のトレース実験を行ったところ,地上部において標識されたトランスゼアチンは検出されなかった.これらの結果から,根に投与したトランスゼアチンは代謝をうけずに地上部へと長距離輸送されることが示された.\r\n 根から長距離輸送されたトランスゼアチンが地上部において実際に作用するかどうか検証するため,log七重変異体を穂木,野生型を台木とした接木植物の地上部の表現型を観察した.log七重変異体は野生型と比較して地上部がいちじるしく小さくなることが報告されているが4),この接木植物の地上部の大きさは野生型と差がなかった.また,log七重変異体を穂木,abcg14変異体を台木とした接木植物の地上部の大きさはlog七重変異体と差がなかった.さらに,野生型,log七重変異体,log七重変異体と野生型との接木植物,log七重変異体とabcg14変異体との接木植物の地上部において,トランスゼアチンの内生量およびサイトカイニン応答のマーカー遺伝子の発現量を解析したところ,表現型とのあいだにみかけ上の相関がみられた.以上の結果から,根から地上部へ長距離輸送されたトランスゼアチンは地上部において作用し,地上部の大きさを維持する作用をもつことが示唆された.\r\n\r\n3.根から長距離輸送されたトランスゼアチンとトランスゼアチンリボシドの地上部の成長の制御における役割は異なる\r\n 道管を長距離輸送されたトランスゼアチンとその前駆体であるトランスゼアチンリボシドのサイトカイニンとしての作用に違いがあるかどうか検討するため,2種類のサイトカイニン生合成系遺伝子の変異体を穂木,野生型を台木とした接木植物を作出し,地上部の表現型を詳細に解析した.一方の接木植物はlog七重変異体を穂木とした接木植物で,地上部におけるトランスゼアチンの生合成能は欠損するが,根から長距離輸送されたトランスゼアチンは地上部において作用する.また,サイトカイニン活性化経路を欠損することから,根から長距離輸送されたトランスゼアチンリボシドは地上部にて活性化されず地上部においては作用しない.もう一方の接木植物はサイトカイニン前駆体生合成酵素IPTの変異体であるipt三重変異体を穂木とした接木植物である.IPTはサイトカイニン生合成の初期反応を触媒することから,ipt三重変異体においてはトランスゼアチンおよびトランスゼアチンリボシドの生合成はいちじるしく低下し地上部が小さくなる6).しかし,外部からトランスゼアチンリボシドなどサイトカイニン前駆体を投与すればLOGによりこれを活性化することができ,地上部の成長は回復する.したがって,この接木植物においては,地上部におけるトランスゼアチンの生合成が低下する一方,根から長距離輸送されたトランスゼアチンとトランスゼアチンリボシドの両方が地上部において作用する.\r\n この2つの接木植物の地上部を観察したところ,みかけの大きさは野生型と差がなかった.しかし,地上部の大きさについて詳細に解析するといくつか違いがあった.葉の面積はlog七重変異体およびipt三重変異体において小さくなったのに対し,野生型,log七重変異体を穂木とした接木植物,ipt三重変異体を穂木とした接木植物のあいだには有意な差はなかった.一方,葉の形成の速度を比較すると,野生型とipt三重変異体を穂木とした接木植物とのあいだに差はなかったが,log七重変異体,ipt三重変異体,log七重変異体を穂木とした接木植物においては野生型に比べ有意に遅くなった.葉の形成の速度にかかわると予想される茎頂分裂組織を観察すると,その大きさは葉の形成の速度の関係と一致した.さらに,地上部におけるトランスゼアチンの内生量およびサイトカイニン応答のマーカー遺伝子の発現量を解析すると,log七重変異体およびipt三重変異体においては低下していたのに対し,野生型,log七重変異体を穂木とした接木植物,ipt三重変異体を穂木とした接木植物のあいだには差がなかった.したがって,葉の形成の速度についてlog七重変異体を穂木とした接木植物とipt三重変異体を穂木とした接木植物の違いを生じる原因は,単なるトランスゼアチンの存在量ではないことが示された.これらの結果から,根から長距離輸送されたトランスゼアチンは葉の面積の維持にのみ作用する一方,根から長距離輸送されたトランスゼアチンリボシドは葉の面積の維持にくわえ葉の形成の速度にも作用することが示唆された(図2).したがって,根から長距離輸送されたトランスゼアチンとその前駆体であるトランスゼアチンリボシドは地上部の成長の制御において異なる役割を担うことが示された.\r\n\r\n4.長距離輸送されるトランスゼアチンとトランスゼアチンリボシドとの比は窒素栄養の環境により異なる\r\n 根から地上部へ長距離輸送されるトランスゼアチンとトランスゼアチンリボシドは異なる成長形質を制御することが示されたが,その生物学的な意義はなんであろうか.この問いに答えるため,窒素栄養の環境が変動する状況にて道管液におけるトランスゼアチンおよびトランスゼアチンリボシドの濃度を解析した.その結果,窒素栄養の添加に対するトランスゼアチンリボシドの濃度の上昇は,トランスゼアチンの濃度の上昇に比べ数倍も高かった.したがって,長距離輸送されるトランスゼアチンとトランスゼアチンリボシドとの比は根圏の栄養環境により異なることが示された.このことから,サイトカイニンによる葉の面積および葉の形成の速度の促進作用は必ずしも同等ではなく,窒素の富栄養条件においては新たな葉の形成をより促進させることが示唆された.\r\n\r\nおわりに\r\n これまでに,筆者らの研究グループは,サイトカイニンの作用は量の変化だけでなく,サイトカイニン分子の側鎖の修飾による質的な変化によっても制御されることを明らかにしてきた7)(新着論文レビュー でも掲載).今回の研究において,さらに,同じ側鎖の構造をもつサイトカイニン分子でも,輸送のときの形態により異なる作用をもつことが明らかにされた.長距離輸送される植物ホルモンとしては,ほかにもオーキシン,アブシジン酸,ジベレリン,ストリゴラクトンなどが知られているが,その輸送の形態と作用との関係についてあまりくわしくは調べられていない.今回の発見が,長距離輸送される植物ホルモンの複雑な形質制御にはたす役割の分子レベルでの理解につながることを期待したい.\r\n\r\n〈文 献〉\r\n 1) Kieber, J. J. \u0026 Schaller, G. E.: Cytokinins. Arabidopsis Book, 12, e0168 (2014)\r\n 2) Sakakibara, H.: Cytokinins: activity, biosynthesis, and translocation. Annu. Rev. Plant Biol., 57, 431-449 (2006)\r\n 3) Takei, K., Sakakibara, H., Taniguchi, M. et al.: Nitrogen-dependent accumulation of cytokinins in root and the translocation to leaf: implication of cytokinin species that induces gene expression of maize response regulator. Plant Cell Physiol., 42, 85-93 (2001)\r\n 4) Tokunaga, H., Kojima, M., Kuroha, T. et al.: Arabidopsis lonely guy (LOG) multiple mutants reveal a central role of the LOG-dependent pathway in cytokinin activation. Plant J., 69, 355-365 (2012)\r\n 5) Ko, D., Kang, J., Kiba, T. et al.: Arabidopsis ABCG14 is essential for the root-to-shoot translocation of cytokinin. Proc. Natl. Acad. Sci. USA, 111, 7150-7155 (2014)\r\n 6) Miyawaki, K., Tarkowski, P., Matsumoto-Kitano, M. et al.: Roles of Arabidopsis ATP/ADP isopentenyltransferases and tRNA isopentenyltransferases in cytokinin biosynthesis. Proc. Natl. Acad. Sci. USA, 103, 16598-16603 (2006)\r\n 7) Kiba, T., Takei, K., Kojima, M. et al.: Side-chain modification of cytokinins controls shoot growth in Arabidopsis. Dev. Cell. 27, 452-461 (2014) [新着論文レビュー]\r\n\r\n〈著者プロフィール〉\r\n大薄 麻未(Asami Osugi)\r\n略歴:2012年 東京大学大学院新領域創成科学研究科 修了,同年 理化学研究所環境資源科学研究センター 特別研究員,2016年 名古屋大学大学院生命農学研究科 研究員を経て,2017年よりカナダToronto大学Postdoctoral Fellow.\r\n関心事:植物の器官のあいだ,および,生物のあいだの情報の伝達.\r\n\r\n榊原 均(Hitoshi Sakakibara)\r\n名古屋大学大学院生命農学研究科 教授,理化学研究所環境資源科学研究センター グループディレクター 兼任.\r\n研究室URL:https://www.agr.nagoya-u.ac.jp/~biochem/\r\n\r\n〈図説明〉\r\n図1 接木技術を用いた分子の長距離輸送およびその作用の検証\r\n(a)タグを付加した形質転換体を用いた長距離輸送の検証.\r\n(b)変異体を用いた内在性の分子の長距離輸送の検証.\r\n図2 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