FirstAuthors:8373 JSONTXT

{"target":"http://pubannotation.org/docs/sourcedb/FirstAuthors/sourceid/8373","sourcedb":"FirstAuthors","sourceid":"8373","text":"〈タイトル〉Polycomb群タンパク質は転写の活性化にも寄与する\r\n\r\n〈著 者〉近藤 隆・古関明彦\r\n〈著者所属〉理化学研究所統合生命医科学研究センター 免疫器官形成研究グループ\r\n〈著者email〉tkondo@rcai.riken.jp（近藤 隆）\r\n\r\n〈対象論文〉\r\nPolycomb potentiates Meis2 activation in midbrain by mediating interaction of the promoter with a tissue-specific enhancer.\r\nTakashi Kondo, Kyoichi Isono, Kaori Kondo, Takaho A. Endo, Shigeyoshi Itohara, Miguel Vidal, Haruhiko Koseki\r\nDevelopmental Cell, 28, 94-101 (2014)\r\n\r\n〈要 約〉\r\n　Polycomb群タンパク質は転写抑制性のクロマチンタンパク質であり，ヒストンの修飾，ゲノムDNA領域の凝縮などの活性をもつことが知られている．その標的遺伝子の多くは発生における制御遺伝子であり，動物の形態形成においてその発現を複雑に変化させることが知られている．この研究において，Polycomb群タンパク質はDNA領域のあいだの相互作用を仲介する活性をもつことに着目し，異なる遺伝子発現の状態における，複数の転写制御領域のあいだの相互作用，および，染色体における立体配置の変化を解析することにより，Polycomb群タンパク質が転写の抑制のみならず，活性化にも寄与することを示した．\r\n\r\nはじめに\r\n　Polycomb群タンパク質はショウジョウバエにおいて発見された一群の転写抑制性のクロマチンタンパク質である．大きく分けてPolycomb抑制複合体1（PRC1）およびPolycomb抑制複合体2（PRC2）の2つの複合体を形成し，PRC1はヒストンH2Aの119番目のLysのユビキチン化活性，PRC2はヒストンH3の27番目のLysのトリメチル化活性をもち，転写を抑制する機能をもつ．一方で，近年，Polycomb群タンパク質は染色体における遺伝子座のあいだの相互作用にも関与していることが知られるようになってきた1-4)．また，Polycomb群タンパク質の標的遺伝子には発生における制御遺伝子が多く含まれることが知られている5)．動物の発生においてこれらの制御遺伝子の発現は抑制と活性化をくり返すことが知られており，Polycomb群タンパク質による転写の抑制は，遺伝子発現の活性化の機構とも不可分ではないと考えられる．Polycomb群タンパク質をノックアウトしたマウスの解析において遺伝子発現の活性化に問題のあることが知られていたにもかかわらず6,7)，これまで，Polycomb群タンパク質の転写の活性化に対する寄与については検討されていなかった．筆者らは，マウスのMeis2遺伝子をモデルとして，その転写の抑制あるいは活性化の過程におけるPolycomb群タンパク質の寄与について検討することにより，Polycomb群タンパク質による標的遺伝子の転写の活性化にPolycomb群タンパク質による染色体の立体配置の制御が関与していることを示した．\r\n\r\n1．Meis2遺伝子の転写制御におけるPolycomb群タンパク質の影響\r\n　Meis2遺伝子はHox遺伝子群の補因子であることが知られており8)，発生において時期および組織に特異的に発現することが知られている．とくに，胎生9日目において前脳および中脳での顕著に発現し，また，肢芽では発現が抑制されていることが知られている．Polycomb抑制複合体1の構成タンパク質のひとつであり，そのヒストンへのユビキチン化活性における活性サブユニットであるRING1Bに対する抗体を用いて，ES細胞あるいはさまざまな組織についてクロマチン免疫沈降-シークエンシング（ChIP-seq）法により解析した．その結果，Meis2遺伝子のプロモーター領域および3'側の転写終結領域において著明なピークがみられ，Meis2遺伝子はPolycomb群タンパク質により転写制御をうけていることが示唆された．そこで，RING1のダブルノックアウトマウス（RING1には，ほぼ同様の活性をもつRING1AおよびRING1Bの2つのアイソザイムが存在し，この研究においては，その両方をノックアウトしたダブルノックアウトマウスを使用した）を観察したところ，Meis2遺伝子は胎生11日目の胚における発現パターンをほとんど失い，胴体における過剰な発現がみられたと同時に，本来，前脳および中脳においてみられるはずの強い発現がみられなかった．\r\n\r\n2．Meis2遺伝子の転写を抑制したときの染色体の高次構造\r\n　Meis2遺伝子は野生型のマウスの胎生11日目の胚において肢芽などで発現が抑制されている．それら抑制されている組織におけるクロマチン免疫沈降（ChIP）法による解析により，Meis2遺伝子のプロモーター領域にはRING1Bが蓄積していることが判明している．同時に，そこから200 kb離れたMeis2遺伝子の3'側の転写終結領域におけるRING1Bの結合も顕著に観察される．そこで，蛍光in situハイブリダイゼーション（FISH）法によりこれら2つのDNA領域の核における局在を検出したところ，共局在していることが見い出された．また，タンパク質とDNAを同時に検出する免疫FISH法により，RING1Bも同時に局在していることが見い出された．RING1ダブルノックアウトマウスではMeis2遺伝子の転写の抑制が失われたのと同時に，プロモーター領域と3'側の転写終結領域との相互作用も失われた．これらのことから，RING1Bに依存的に2つのDNA領域は相互作用しており，その状態でMeis2遺伝子に転写の抑制が起こることが示された．\r\n\r\n3．中脳が発生するときのMeis2遺伝子の転写制御\r\n　Meis2遺伝子は中脳が発生するとき発現するため，トランスジェニックマウスなどを用いて，そのとき機能しているエンハンサー領域を同定した．FISH法による解析により，そのエンハンサー領域は中脳において特異的にMeis2遺伝子のプロモーター領域と相互作用していることが見い出された．おもしろいことに，RING1ダブルノックアウトマウスでは中脳におけるMeis2遺伝子の強い発現が失われると同時に，このエンハンサー領域とプロモーター領域との相互作用も失われたことが見い出された．\r\n　Meis2遺伝子の発現について，時系列をおってより詳細に調べた．中脳における発現は胎生9日目の胚の22体節のときから検出されはじめたので，胎生6日目の胚の胚形成予定領域（エピブラスト），16体節胚から20体節胚の中脳予定領域，22体節胚から26体節胚の中脳を用いて，Meis2遺伝子のプロモーター領域，エンハンサー領域，3'側の転写終結領域の3つのDNA領域の立体配置について検討した．胎生6日目の胚ではプロモーター領域と3'側の転写終結領域とが相互作用しておりエンハンサー領域は共存していなかった．22体節胚から26体節胚ではプロモーター領域と相互作用しているのはエンハンサー領域に換わっており，3'側の転写終結領域は共存していなかった．16体節胚から20体節胚ではこれら3つのDNA領域はすべて集合していることが見い出され，一時的な中間段階の存在することが示された．RING1ダブルノックアウトマウスでは16体節胚から20体節胚において3つのDNA領域が共局在する現象が見い出されなかったため，この3つのDNA領域の集合はRING1に依存性であると考えられた．\r\n\r\nおわりに\r\n　この研究において，転写の活性化は染色体における転写制御領域のあいだの空間的な配置，とくに，プロモーター領域との物理的な接触あるいは相互作用が重要であることが示唆されたのと同時に，Polycomb群タンパク質のひとつであるRING1がその空間的な配置の決定に重要であることが示された．これまで，Polycomb群タンパク質は転写の抑制の維持に重要であると考えられてきたが，この研究において，エンハンサー領域とプロモーター領域とのあいだの相互作用をも制御していることが示され，Polycomb群タンパク質は転写の活性化においても重要であることが示された（図1）．Polycomb群タンパク質の標的遺伝子の多くは発生において重要なはたらきをもつことが知られており，それらの制御遺伝子の発現は生体の活動において活性化と抑制をくり返すことが知られている．このことから，Polycomb群タンパク質は転写を再活性化する可能性を維持しながら転写を抑制することが主たる役割であることが予想される．この研究において明らかにされた機構は，Polycomb群タンパク質による転写の制御における重要な側面であると考えられた．\r\n\r\n〈文 献〉\r\n 1) Eskeland, R., Leeb, M., Grimes, G. R. et al.: Ring1B compacts chromatin structure and represses gene expression independent of histone ubiquitination. Mol. Cell, 38, 452-464 (2010)\r\n 2) Bantignies, F., Roure, V., Comet, I. et al.: Polycomb-dependent regulatory contacts between distant Hox loci in Drosophila. Cell, 144, 214-226 (2011)\r\n 3) Endoh, M., Endo, T. A., Endoh, T. et al.: Histone H2A mono-ubiquitinaton is a crucial step to mediate PRC1-dependent repression of developmental genes to maintain ES cell identity. PLoS Genet., 8, e1002774 (2012)\r\n 4) Isono, K., Endo, T. A., Ku, M. et al.: SAM domain polymerization links subnuclear clustering of PRC1 to gene silencing. Dev. Cell, 26, 565-577 (2013) [新着論文レビュー]\r\n 5) Tanay, A., O'Donnell, A. H., Damelin, M. et al.: Hyperconserved CpG domains underlie Polycomb-binding sites. Proc. Natl. Acad. Sci. USA, 104, 5521-5526 (2007)\r\n 6) Akasaka, T., van Lohuizen, M., van der Lugt, N. et al.: Mice doubly deficient for Polycomb Group genes Mel18 and Bmi1 reveal synergy and requirement for maintenance but not initiation of Hox gene expression. Development, 128, 1587-1597 (2001)\r\n 7) de Graaff, W., Tomotsune, D., Oosterveen, T. et al.: Randomly inserted and targeted Hox/reporter fusions transcriptionally silenced in Polycomb mutants. Proc. Natl. Acad. Sci. USA, 100, 13362-13367 (2003)\r\n 8) Rieckhof, G. E., Casares, F., Ryoo, H. D. et al.: Nuclear translocation of extradenticle requires homothorax, which encodes an extradenticle-related homeodomain protein. Cell, 91, 171-183 (1997)\r\n\r\n〈著者プロフィール〉\r\n近藤 隆（Takashi Kondo）\r\n略歴：1992年 東京大学大学院医学研究科博士課程 修了，1994年 癌研究会癌研究所 嘱託研究員，2000年 スイスGeneva大学 博士研究員，2010年 理化学研究所脳科学総合研究センター ユニットリーダー，2013年 理化学研究所統合医科学研究センター 研究員を経て，同年より神奈川科学技術アカデミー サブリーダー．\r\n研究テーマ：生命の活動における遺伝子の発現制御と，核あるいは染色体の高次構造のエピジェネティックな相関．\r\n抱負：平穏な日々を過ごすこと．\r\n\r\n古関 明彦（Haruhiko Koseki）\r\n理化学研究所統合医科学研究センター グループディレクター．\r\n研究室URL：http://web.rcai.riken.jp/en/labo/genetics/\r\n\r\n〈図説明〉\r\n図1　Meis2遺伝子の転写制御におけるモデル\r\n（a）野生型のマウス．Meis2遺伝子の転写が抑制されているときには，プロモーター領域と3'側の転写終結領域にあるRING1B結合配列がPolycomb群タンパク質の活性により相互作用しており，転写開始の準備状態ではPolycomb群タンパク質に依存してエンハンサー領域がそこにくわわるかたちでこれら3つのDNA領域の共存状態が一時的に形成される．それにより，エンハンサー領域とプロモーター領域との相互作用が確立され，エンハンサー領域の作用によりRING1B結合配列およびRING1Bがプロモーター領域からはずれ，Meis2遺伝子の転写が活性化される．\r\n（b）RING1ダブルノックアウトマウス．Polycomb群タンパク質の非存在化ではこれらの過程が生ずることなく，転写の抑制も活性化も起こらないため，Meis2遺伝子は弱い発現を継続する．\r\n","tracks":[{"project":"FA_Top107-forWeb","denotations":[{"id":"T1","span":{"begin":6,"end":20},"obj":"polycomb"},{"id":"T2","span":{"begin":424,"end":438},"obj":"polycomb"},{"id":"T3","span":{"begin":565,"end":579},"obj":"polycomb"},{"id":"T4","span":{"begin":675,"end":689},"obj":"polycomb"},{"id":"T5","span":{"begin":728,"end":742},"obj":"polycomb"},{"id":"T6","span":{"begin":929,"end":943},"obj":"polycomb"},{"id":"T7","span":{"begin":995,"end":1009},"obj":"polycomb"},{"id":"T8","span":{"begin":1090,"end":1104},"obj":"polycomb"},{"id":"T9","span":{"begin":1142,"end":1156},"obj":"polycomb"},{"id":"T10","span":{"begin":1216,"end":1230},"obj":"polycomb"},{"id":"T11","span":{"begin":1305,"end":1319},"obj":"polycomb"},{"id":"T12","span":{"begin":1336,"end":1350},"obj":"polycomb"},{"id":"T13","span":{"begin":1366,"end":1380},"obj":"polycomb"},{"id":"T14","span":{"begin":1431,"end":1445},"obj":"polycomb"},{"id":"T15","span":{"begin":1767,"end":1781},"obj":"polycomb"},{"id":"T16","span":{"begin":3370,"end":3384},"obj":"polycomb"},{"id":"T17","span":{"begin":3427,"end":3441},"obj":"polycomb"},{"id":"T18","span":{"begin":3517,"end":3531},"obj":"polycomb"},{"id":"T19","span":{"begin":3560,"end":3574},"obj":"polycomb"},{"id":"T20","span":{"begin":3660,"end":3674},"obj":"polycomb"},{"id":"T21","span":{"begin":3740,"end":3754},"obj":"polycomb"},{"id":"T22","span":{"begin":5820,"end":5834},"obj":"polycomb"},{"id":"T23","span":{"begin":5860,"end":5874},"obj":"polycomb"},{"id":"T24","span":{"begin":6051,"end":6065},"obj":"polycomb"},{"id":"T25","span":{"begin":445,"end":455},"obj":"chromatin"},{"id":"T26","span":{"begin":769,"end":779},"obj":"chromatin"},{"id":"T27","span":{"begin":849,"end":856},"obj":"Histone 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