FA@AikoHIRAKI:8776_ JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/FA@AikoHIRAKI/sourceid/8776_","sourcedb":"FA@AikoHIRAKI","sourceid":"8776_","text":"〈タイトル〉アブシジン受容体のアンタゴニストの合理的な創出\r\n\r\n〈著 者〉竹内 純1・岡本昌憲2・轟 泰司1\r\n〈著者所属〉1静岡大学大学院農学研究科 応用生物化学専攻天然物有機化学研究室，2鳥取大学乾燥地研究センター 生物生産部門植物分子生物化学研究室\r\n〈著者email〉f5144030@ipc.shizuoka.ac.jp（竹内 純），okamo@alrc.tottori-u.ac.jp（岡本昌憲），aytodor@ipc.shizuoka.ac.jp（轟 泰司）\r\n\r\n〈対象論文〉\r\nDesigned abscisic acid analogs as antagonists of PYL-PP2C receptor interactions.\r\nJun Takeuchi, Masanori Okamoto, Tomonori Akiyama, Takuya Muto, Shunsuke Yajima, Masayuki Sue, Mitsunori Seo, Yuri Kanno, Tsunashi Kamo, Akira Endo, Eiji Nambara, Nobuhiro Hirai, Toshiyuki Ohnishi, Sean R. Cutler , Yasushi Todoroki\r\nNature Chemical Biology, 10, 477-482 (2014)\r\n\r\n〈要 約〉\r\n　アブシジン酸は種子の休眠や環境ストレス応答において重要な役割を担う植物ホルモンである．アブシジン酸受容体はアブシジン酸との結合によりその配座が変化し，プロテインホスファターゼであるPP2Cと結合してその酵素活性を阻害することにより，それ以降のシグナル伝達を活性化する．筆者らは，アブシジン酸と同様にアブシジン酸受容体に高い親和性をもちながら，アブシジン酸受容体とPP2Cとのあいだの相互作用を阻害するアブシジン酸受容体のアンタゴニストを，アブシジン酸受容体とアブシジン酸との複合体の結晶構造に基づき設計した．アブシジン酸受容体-アブシジン酸複合体にはアブシジン酸の3'-CHの方向に小さなトンネルが存在し，これはPP2Cとの結合表面へとつうじていた．そこで，このトンネルをとおってアブシジン酸受容体の外側にアルキル鎖を突出する化合物はアブシジン酸受容体とPP2Cとのあいだの相互作用を妨害してアブシジン酸受容体のアンタゴニストとして機能する可能性が高いと考え，アブシジン酸の3'位に硫黄を介して直鎖アルキル鎖を導入した化合物を合成した．種子の発芽および乾燥耐性の誘導，アブシジン酸応答性遺伝子の発現，アブシジン酸受容体-アブシジン酸複合体によるPP2Cの活性阻害に対する効果を検証することにより，この化合物は直鎖アルキル鎖の長さによりアゴニストあるいはアンタゴニストとして機能することが確認された．さらに，アブシジン酸受容体との複合体の結晶構造解析により，この化合物はトンネルから突出したヘキシル鎖がアブシジン酸受容体とPP2Cとのあいだの相互作用を妨害することにより，アブシジン酸受容体のアンタゴニストとして機能することが明らかにされた．\r\n\r\nはじめに\r\n　植物が乾燥ストレスなどを受けた際に合成されたアブシジン酸は，標的となる器官に輸送されたのち，STARTドメインスーパーファミリーと構造的によく類似した細胞質受容体であるアブシジン酸受容体と結合してさまざまな生理作用を誘導する1,2)．シロイヌナズナのアブシジン酸受容体はアブシジン酸に対する依存性の相違から二量体型受容体と単量体型受容体とに分類される3,4)．二量体型受容体はアブシジン酸との結合により単量体に解離し，ゲート開口型配座からゲート閉鎖型配座へと変化して，アブシジン酸シグナル伝達系における負の制御タンパク質であるプロテインホスファターゼPP2Cとの結合表面を形成する．そののち，アブシジン酸受容体-アブシジン酸複合体はPP2Cと結合してその活性を阻害する5,6)．これにより，PP2Cにより不活性化されていたプロテインキナーゼSnRK2は自己リン酸化によりその活性を回復し7)，イオンチャネルや転写因子を活性化してアブシジン酸応答を誘導する8)（図1）．一方，単量体型受容体はアブシジン酸が存在しなくてもゲート開口型配座とゲート閉鎖型配座の平衡状態にあるため，弱いながらPP2Cの活性を阻害することができる2-4)．これまでの研究から，アブシジン酸応答にはどちらのタイプのアブシジン酸受容体も関与していることがわかっているが，単量体型受容体によるアブシジン酸に非依存的なPP2Cの阻害の生理的な意義についてはいまだよくわかっていない．\r\n　これまで，アブシジン酸受容体のアゴニストに関しては多く報告されているが9,10)，アブシジン酸受容体のアンタゴニストに関する研究はほとんど行われていない．アブシジン酸受容体アンタゴニストはシグナル伝達の起点に位置する受容体をターゲットとするため，アブシジン酸応答を網羅的に阻害することが可能であり，遺伝子の解析の進んでいない非モデル植物においてもアブシジン酸の多様な生理作用を解析する際の有用な化学ツールになることが期待される．また，アブシジン酸は種子の発芽の阻害や成長の制御にも関与することから，アブシジン酸受容体アンタゴニストは植物の成長の調節剤として農業分野への応用も期待される．\r\n　アブシジン酸受容体のアンタゴニストを合理的に創出するためには，アブシジン酸受容体に対する高い選択性とアブシジン酸受容体とPP2Cとのあいだの相互作用の阻害とを同時に実現するデザインが必要である．そこで，この研究では，アロステリック制御因子として作用するアブシジン酸の構造的な特徴およびアブシジン酸受容体の作用機構に着目し，アブシジン酸受容体-アブシジン酸複合体の結晶構造にもとづいてアブシジン酸受容体アンタゴニストを設計し，その効果をin vitroおよびin vivoにおいて検証した．\r\n\r\n1．アブシジン酸受容体のアンタゴニストの分子設計\r\n　アブシジン酸受容体-アブシジン酸複合体において，アブシジン酸の側鎖のカルボン酸はアブシジン酸受容体のもつポケットの奥に位置しLysの側鎖と塩橋を形成して強く固定されている．したがって，アブシジン酸の側鎖の修飾はアブシジン酸受容体との親和性をいちじるしく損ねると考えられ，構造活性相関もこれを強く支持していた．一方，アブシジン酸のシクロへキセノン環の部分はアブシジン酸受容体のもつポケットの入口の付近に位置し，疎水性相互作用によるアブシジン酸受容体におけるゲートの閉鎖の誘導に関与する．アブシジン酸受容体-アブシジン酸複合体において，アブシジン酸はアブシジン酸受容体のもつポケットに完全に包み込まれているが，アブシジン酸の3'-CHの方向には配座の変化により生じた小さな疎水性のトンネルが存在し，アブシジン酸受容体の表面にあるPP2C結合部位へとつうじている．また，アブシジン酸の3'位の修飾体はアブシジン酸と同じ程度の活性を保持していることから，アブシジン酸の3'位へのアルキル鎖の導入によりアゴニストをアンタゴニストへと転換できるかもしれないと考えた．そこで，アブシジン酸のシクロへキセノン環の二重結合をエポキシ化したのち，アルキルチオレートによるエポキシ開環と，それにつづくin situ脱水反応により，化合物ASn（n = 2〜12）を合成した．アブシジン酸受容体とASnとの複合体のモデルでは，nが4のときアルキル鎖の長さがアブシジン酸受容体のもつトンネルの長さとほぼ一致したため，ASnは，nが4以下でアゴニスト，nが4以上でアンタゴニストとして機能すると予測した．\r\n\r\n2．植物に対する生理活性\r\n　モデル植物であるシロイヌナズナに対するASnの生理活性について調べた．ASnをそれぞれ単独で処理したところ，AS2〜AS4はアブシジン酸と同様に種子の発芽の阻害および実生の初期生育の阻害を示し，活性の強いほうからAS2，アブシジン酸，AS3，AS4の順であった．一方，AS5〜AS12に関してはアブシジン酸様の阻害効果は観察されなかった．アブシジン酸との共処理によりASnのアンタゴニスト活性を評価したところ，AS2〜AS4はアブシジン酸の効果を増強し，AS5〜AS12はアブシジン酸による発芽の阻害および初期生育の阻害の効果を抑制した．ASnはレタスの種子の発芽および初期生育に対してもシロイヌナズナの場合と同様の効果を示した．また，ダイコンの実生を用いて乾燥耐性の誘導に対するASnの効果を評価した．この試験では，乾燥処理により増加する内生のアブシジン酸に対する拮抗作用を評価するためにASnを単独で処理した．その結果，AS2はアブシジン酸と同様に乾燥耐性を増強する効果を示した一方，AS6は乾燥処理による内生アブシジン酸の増加に拮抗して乾燥耐性を減弱する効果を示した．\r\n　アブシジン酸に応答性の遺伝子発現に対するASnの効果を，定量RT-PCR法およびアブシジン酸応答性遺伝子のプロモーターにレポーター遺伝子を導入したトランスジェニックシロイヌナズナを用いて調べた．その結果，AS2はアブシジン酸よりやや強く，濃度に依存した遺伝子発現を誘導し，また，アブシジン酸よりも強いレポーター遺伝子の発現を示した．AS4による遺伝子発現の誘導は定量RT-PCR法ではほとんど検出できなかったが，レポーター遺伝子アッセイにおいては根においてわずかにレポーター遺伝子の発現がみられた．AS6ではどちらの試験においても遺伝子発現の誘導はみられなかった．アブシジン酸との共処理においては，AS2はアブシジン酸による遺伝子発現の誘導を強め，反対に，AS6およびAS4はアブシジン酸による遺伝子発現の誘導を明確に抑制し，その抑制効果はAS6においてより顕著であった．さらに，AS6はマンニトールの処理による浸透圧ストレスにより誘導される内生のアブシジン酸によるレポーター遺伝子の発現も抑制した．\r\n　以上の結果から，AS2はアブシジン酸受容体のアゴニスト，AS6はアンタゴニスト，そして，AS4はアゴニストとアンタゴニストの中間的な機能をもつことが示唆された．\r\n\r\n3．アブシジン酸受容体とPP2Cとのあいだの相互作用に対する生化学的な活性\r\n　アブシジン酸受容体においてトンネルを構成するアミノ酸残基はアブシジン酸受容体のアイソフォームのあいだで高度に保存されていたことから，ASnはアイソフォームに関係なく活性を示すと予測された．そこで，アブシジン酸受容体のアイソフォームとして，シロイヌナズナのPYR1，PYL1〜PYL6，PYL8〜PYL11（PYL7とPYL12に関しては機能的なタンパク質が得られなかった）による，PP2Cの脱リン酸化活性の阻害に対するASnの効果について検証した．この試験では，アゴニストはほぼ完全にPP2Cの活性を阻害する一方，パーシャルアゴニストの場合は不完全にしかPP2Cの活性を阻害できない．アンタゴニストはアブシジン酸との共処理においてアブシジン酸によるPP2Cの活性阻害を抑制し脱リン酸化活性を回復させる．ASnをそれぞれ単独で処理したところ，AS2は二量体型受容体であるPYR1およびPYL1〜PYL3に対しては，アブシジン酸と同じ程度のPP2Cの活性阻害を示した．一方，単量体型受容体に対しては，PYL4，PYL5，PYL11についてアブシジン酸よりも弱いPP2Cの活性阻害を示し，PYL6〜PYL10についてはPP2Cの活性阻害をほとんど誘導しなかった．AS4は二量体型受容体に対しては不完全であるがPP2Cの活性阻害を誘導し，単量体型受容体に対してはPP2Cの活性阻害をまったく誘導しなかった．AS6もAS4と同様に二量体型受容体に対してのみPP2Cの活性阻害を誘導したが，その効果はAS4よりもさらに弱かった．のちに述べるように，AS6はアブシジン酸と同等のアブシジン酸受容体との結合親和性をもつにもかかわらず，PP2Cの活性を50％程度しか阻害できなかったことから，AS6はパーシャルアゴニストであることが示唆された．\r\n　ASnのアンタゴニスト活性を評価するためアブシジン酸との共処理試験を行った．AS4およびAS6は試験したすべてのアブシジン酸受容体に対しアブシジン酸に依存的なPP2Cの活性阻害を抑制し，アブシジン酸受容体のアイソフォームに関係なくアンタゴニストとして機能した．AS6はAS4よりも強いアンタゴニスト活性を示したものの，アブシジン酸に対し20モル当量を処理した場合でもPP2Cの活性を完全に回復させることはできなかった．これは，AS6のパーシャルアゴニスト活性に起因すると考えられた．つまり，AS6はAS6を単独で処理したときのPP2Cの活性値（50％阻害）までしか，アブシジン酸によるPP2Cの活性阻害を回復することができない．\r\n　アブシジン酸受容体-アブシジン酸-PP2C複合体の形成に対するAS6の効果をプルダウンアッセイにより調べた．アブシジン酸との共処理において，AS6はアブシジン酸に依存的なアブシジン酸受容体とPP2Cとのあいだの相互作用を濃度に依存して阻害した．この結果から，AS6はアブシジン酸の誘導するアブシジン酸受容体とPP2Cとのあいだの相互作用を阻害することにより，アブシジン酸受容体のアンタゴニストとして機能することが実証された．\r\n\r\n4．熱力学的な解析および結晶構造解析\r\n　等温滴定型熱量測定によりアブシジン酸受容体とAS6との複合体の形成を熱力学的に解析した．アブシジン酸受容体として，リガンドとの結合の際に二量体の解離の影響のない単量体型受容体であるPYL5およびPYL10を用いてAS6の解離定数を測定したところ，アブシジン酸と同じ程度の結合親和性を示した．AS6はPYL5に対し，アブシジン酸と比較して結合の際の発熱量（エントロピー変化）は大きかったものの，エントロピー的には不利であり，アブシジン酸と同じ程度の親和性を示した．一方，PYL10に対しては真逆で，結合の際の発熱量は減少した反面，エントロピー損失も低減したため，総合的にはアブシジン酸と同じ程度の親和性を示した．AS6によるアブシジン酸受容体の配座の変化においては，アブシジン酸受容体のアイソフォームのあいだでの微妙な違いが影響している可能性がある．\r\n　AS6がアブシジン酸受容体のアンタゴニストであることを分子レベルにおいて決定づけるため，アブシジン酸受容体PYR1とAS6との複合体のX線結晶構造解析を行った．PYR1-AS6複合体の構造はPYR1-アブシジン酸複合体の構造とよく類似しており，AS6はアブシジン酸と同じ配座でPYR1のもつリガンド結合ポケットに結合してゲートの閉鎖を誘導することが確認された．さらに，分子設計の際のアブシジン酸受容体-ASn複合体のモデルと一致して，AS6はPYR1のもつトンネルから突出したヘキシル鎖が立体障害となってアブシジン酸受容体とPP2Cとのあいだの相互作用を妨害することにより，アブシジン酸受容体のアンタゴニストとして機能することが証明された（図2）．\r\n\r\n5．in vivoにおける機能選択性\r\n　これまで述べた実験結果から，ASnの標的がアブシジン酸受容体であることは明確になった．しかし，in vivoにおけるASnの生理活性においてはアブシジン酸受容体以外の因子も関与している可能性が考えられた．ASnは3'位に直鎖アルキル鎖を導入した以外はアブシジン酸の骨格をそのまま保持しているため，潜在的なオフターゲット効果の要因はアブシジン酸代謝酵素とアブシジン酸トランスポーターにほぼ限定された．そこで，それらに対するASnの影響について調べた．\r\n　アブシジン酸は植物においておもにアブシジン酸8'-ヒドロキシラーゼ（CYP707A）により水酸化されたのち最終的に不活性化される．CYP707Aに対するAS2およびAS6の阻害定数を求めたところ，アブシジン酸と比較してCYP707Aに対する親和性は著しく低かった．さらに，AS2あるいはAS6を処理したシロイヌナズナの内生のアブシジン酸の量は無処理の場合と比較して差異がなかったことから，ASnはアブシジン酸の代謝にほとんど影響をあたえないことが示唆された．\r\n　これまで，アブシジン酸トランスポーターとしてはABCトランスポーターであるAtABCG25およびAtABCG40，硝酸トランスポーターファミリーに属するNPF4.6/AIT1が報告されている．このなかで唯一，試験法の確立されていたAIT1を用いて，アブシジン酸の輸送に対するAS6の影響を調べた．その結果，AS6はアブシジン酸に対し10モル当量を処理した場合でもわずかな阻害活性しか示さなかったため，AIT1に対する親和性はアブシジン酸の1/10以下となっている可能性が高いと思われた．\r\n　以上より，ASnはアブシジン酸代謝酵素およびアブシジン酸トランスポーターの強力な阻害剤としては作用しないことが示唆された．\r\n\r\nおわりに\r\n　この研究では，構造活性相関と立体構造情報にもとづいて，アブシジン酸受容体のアンタゴニストを合理的に創出することにはじめて成功した．アブシジン酸のシグナル伝達機構に関する研究はおもにモデル植物において行われており，シロイヌナズナにおいてはその概要が明らかにされている．一方で，分子生物学的な手法や遺伝学的な手法の適用の困難な多くの植物種においては，アブシジン酸の生理作用を分子レベルで理解する研究は遅れている．今回，創出されたアブシジン受容体のアンタゴニストAS6は，シロイヌナズナだけではなくレタスにおける種子の発芽の阻害やカイワレダイコンにおける乾燥耐性の誘導においてもアブシジン酸の生理作用を阻害したことから，非モデル植物におけるアブシジン酸のシグナル伝達機構の解明において有用な化学ツールとなることが期待される．さらに今後，より低濃度で強力に作用する化合物を開発することにより，実用の可能な植物成長の調節剤の創出にもつながることが期待される．\r\n\r\n〈文 献〉\r\n 1) Ma, Y., Szostkiewicz, I., Korte, A. et al.: Regulators of PP2C phosphatase activity function as abscisic acid sensors. Science, 324, 1064-1068 (2009)\r\n 2) Park, S. Y., Fung, P., Nishimura, N. et al.: Abscisic acid inhibits type 2C protein phosphatase via the PYR/PYL family of START proteins. Science, 324, 1068-1071 (2009)\r\n 3) Dupeux, F., Santiago, J., Betz, K. et al.: A thermodynamic switch modulates abscisic acid receptor sensitivity. EMBO J., 30, 4171-4184 (2011)\r\n 4) Hao, Q., Yin, P., Li, W. et al.: The molecular basis of ABA-independent inhibition of PP2Cs by a subclass of PYL proteins. Mol. Cell, 42, 662-672 (2011)\r\n 5) Melcher, K., Ng, L. M., Zhou, X. E. et al.: A gate-latch-lock mechanism for hormone signaling by abscisic acid receptors. Nature, 462, 602-608 (2009)\r\n 6) Miyazono, K., Miyakawa, T., Sawano, Y. et al.: Structural basis of abscisic acid signalling. Nature, 462, 609-614 (2009)\r\n 7) Soon, F. F., Ng, L. M., Zhou, X. E. et al.: Molecular mimicry regulates ABA signaling by SnRK2 kinase and PP2C phosphatases. Science, 335, 85-88 (2012)\r\n 8) Weiner, J. J., Peterson, F. C., Volkman, B. F. et al.: Structural and functional insights into core ABA signaling. Curr. Opin. Plant Biol., 13, 495-502 (2010)\r\n 9) Melcher, K., Xu, Y., Ng, L. M. et al.: Identification and mechanism of ABA receptor antagonism. Nat. Struct. Mol. Biol., 17, 1102-1108 (2010)\r\n10) Todoroki, Y. \u0026 Hirai, N.: Abscisic acid analogs for probing the mechanism of abscisic acid reception and inactivation. Stud. Nat. Prod. Chem., 27, 321-360 (2002)\r\n\r\n〈著者プロフィール〉\r\n竹内 純（Jun Takeuchi）\r\n略歴：静岡大学創造科学技術大学院博士課程 在学中．\r\n研究テーマ：アブシジン酸受容体のアンタゴニストの創出．\r\n関心事：低分子化合物によるタンパク質の機能の制御．\r\n\r\n岡本 昌憲（Masanori Okamoto）\r\n略歴：鳥取大学乾燥地研究センター テニュアトラック助教．\r\n抱負：低分子化合物を利用して植物における未知の生理現象を分子レベルで明らかにしたい．\r\n\r\n轟 泰司（Yasushi Todoroki）\r\n静岡大学大学院農学研究科 教授．\r\n研究室URL：http://www.agr.shizuoka.ac.jp/c/npchem/index.html\r\n\r\n〈図説明〉\r\n図1　アブシジン酸のシグナル伝達機構のモデル\r\n環境ストレスなどを受けた際に合成されたアブシジン酸は，アブシジン酸受容体に結合したのち，アブシジン酸受容体とプロテインホスファターゼPP2Cとのあいだの相互作用をひき起こしてPP2Cのもつ脱リン酸化活性を抑制する．これにより，PP2Cより不活性化されていたプロテインキナーゼSnRK2が自己リン酸化により活性を回復し，ABFなどの転写因子，および，イオンチャネルSLAC1をリン酸化することによりアブシジン酸応答が誘導される．\r\n図2　AS6がアンタゴニストとして機能する機構\r\n（a）PYR1-AS6複合体の結晶構造．\r\n（b）AS6の作用機構．アブシジン酸受容体のもつトンネルから突出したヘキシル鎖が立体障害となってアブシジン酸受容体とPP2Cとのあいだの相互作用を阻害することにより，AS6はアンタゴニストとして機能する．","tracks":[{"project":"FirstAuthor_s_Plants","denotations":[{"id":"T1","span":{"begin":619,"end":625},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T3","span":{"begin":672,"end":678},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T4","span":{"begin":758,"end":764},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T9","span":{"begin":848,"end":854},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T11","span":{"begin":883,"end":889},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T12","span":{"begin":894,"end":900},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T16","span":{"begin":1049,"end":1055},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T19","span":{"begin":1129,"end":1135},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T23","span":{"begin":1372,"end":1378},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T26","span":{"begin":1485,"end":1491},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T27","span":{"begin":1538,"end":1544},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T28","span":{"begin":1584,"end":1590},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T30","span":{"begin":1656,"end":1662},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T32","span":{"begin":1795,"end":1801},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T35","span":{"begin":1930,"end":1936},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T40","span":{"begin":2150,"end":2156},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T41","span":{"begin":2194,"end":2200},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T46","span":{"begin":2399,"end":2405},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T49","span":{"begin":2444,"end":2450},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T53","span":{"begin":2558,"end":2564},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T54","span":{"begin":2572,"end":2578},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T56","span":{"begin":2640,"end":2646},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T58","span":{"begin":2705,"end":2711},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T62","span":{"begin":2800,"end":2806},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T63","span":{"begin":2814,"end":2820},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T65","span":{"begin":2851,"end":2857},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T67","span":{"begin":2930,"end":2936},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T68","span":{"begin":2945,"end":2951},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T69","span":{"begin":2971,"end":2977},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T70","span":{"begin":3029,"end":3035},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T73","span":{"begin":3317,"end":3323},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T74","span":{"begin":3365,"end":3371},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T76","span":{"begin":3424,"end":3430},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T77","span":{"begin":3468,"end":3474},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T78","span":{"begin":3491,"end":3497},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T79","span":{"begin":3628,"end":3634},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T80","span":{"begin":3671,"end":3677},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T81","span":{"begin":3712,"end":3718},"obj":"https://jglobal.jst.go.jp/detail?JGLOBAL_ID=200907018816139453"},{"id":"T82","span":{"begin":3745,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