{"created":"2023-05-15T10:01:16.299040+00:00","id":5789,"links":{},"metadata":{"_buckets":{"deposit":"60511c44-e118-440b-ae97-ea72c650c476"},"_deposit":{"created_by":5,"id":"5789","owner":"5","owners":[5],"pid":{"revision_id":0,"type":"depid","value":"5789"},"status":"published"},"_oai":{"id":"oai:miyazaki-u.repo.nii.ac.jp:00005789","sets":["72","72:38"]},"author_link":["4839","23623"],"control_number":"5789","item_10010_alternative_title_1":{"attribute_name":"その他（別言語等）のタイトル","attribute_value_mlt":[{"subitem_alternative_title":"高等真核生物が新たな環境変化に適応するために、多世代を通してエピジェネティック修飾による転写因子やクロマチン修飾酵素遺伝子などの5ʼ‑近傍上流のクロマチン構造の可塑性を不可逆的に創出することで、プログラム化されていない新しい細胞機能を獲得するバイオ・システム","subitem_alternative_title_language":"ja"}]},"item_10010_description_5":{"attribute_name":"抄録","attribute_value_mlt":[{"subitem_description":"本総説は多世代を通してクロマチン構造の可塑性を不可逆的に創出することによって、プログラム化されていない新しい細胞機能を獲得する高等真核生物の環境適応能力に関するレビューである。最初に、具体的な例として、ニワトリのHDAC2（-/-） DT40 mutants でHDAC2 遺伝子の欠損に起因して劇的に蓄積したIgM H-chain とL-chain が長期の連続培養期間に劇的に減少する多彩な方法のメカニズムを提示する。引き続いて、これらの具体例に基づいて、異常で不快で不利益な環境変化に自らを適応させるために、高等真核生物がプログラム化されていない新たな細胞機能を獲得する能力に関する一般的な仮説的概念を提示する。ニワトリのDT40 野生株では、HDAC2 はスーパーバイザーとして、Pax5, Aiolos, EBF1, OBF1 とIkaros, E2A などの転写因子の遺伝子発現を逆方向（促進的と抑制的）に制御することを介して、IgM H-chain とL-chainの遺伝子発現を間接的に制御する。HDAC2 遺伝子の欠損に伴って、IgM H-chain とL-chain のmRNAs と蛋白質の劇的な蓄積は急速に誘導される。その後、これらの蓄積したmRNAs と蛋白質の量は、同一条件下での単純な連続培養期間に、全てのHDAC2（-/-） individual clones でほぼ同じ変化パターンで劇的に減少する。一方、Pax5, Aiolos, EBF1, OBF1 の遺伝子発現は、連続培養期間に、個々のHDAC2（-/-） individual clones で著しく異なるパターンで変化する。培養の後期ステージでは、IgM H-chain とL-chain の遺伝子発現に関して、調べた6個のHDAC2（-/-） individualclones は少なくともOBF1-依存性、Pax5, Aiolos-依存性、Pax5, Aiolos, EBF1-依存性の3 つの異なるタイプに分類分け出来る。個々のHDAC2（-/-） individual clones におけるPax5, Aiolos, EBF1,OBF1 の遺伝子発現の複雑な変化は、連続培養期間のこれら遺伝子の5'-近傍上流クロマチン領域でのヒストンH3 の特定Lys 残基のアセチル化及び脱アセチル化レベルの多種多様な変化に基づいた多様な不可逆的なクロマチン・コンフォメーション（構造）変化に起因している。これらの結果に基づいて、高等真核生物が、多世代を通して、エピジェネティック修飾によるクロマチン構造の可塑性の不可逆的創出を介して、プログラム化されていない新しい細胞機能を獲得するバイオ・システムを、クロマチン・コンフォメーション・チェンジ・コード説 （chromatin conformation change code theory: 4C theory）と命名した。\n 高等真核生物の生命維持と細胞タイプの決定にとって最も基本的で重要な方法の1 つと考えられる4C theory の具体的な内容は次の通りである。1） 高等真核生物の体細胞は、その生涯で初めて新しい環境変化に遭遇した時、その変化に適応するかまたは克服するために、プログラム化されていない新規の細胞機能を獲得するための柔軟で順応的で多能的な能力を有している。2） 細胞のこの柔軟で順応的な多能性は基本的にはクロマチン構造の柔軟性・順応性・多能性に基づいている。3） 体細胞は、多世代（多数の細胞分裂）を通して、特定の転写因子やクロマチン修飾酵素の遺伝子の5'-近傍上流域のクロマチン構造の可塑性を不可逆的に創出することによって、環境変化に適応または排除するためのプログラム化されていない新しい細胞機能を獲得する能力を有す。4） クロマチン構造（緩いフォームから硬いフォームまたはその逆）の可塑性は、多世代を通したエピジェネティック修飾によるクロマチン・コンフォメーション・チェンジに基づいて、連続的・不可逆的に創出される。5） 同じタイプの個々の細胞におけるクロマチン構造の可塑性の多様性は、環境変化への自発的で不均衡な対応によって引き起され、その後の多世代を通した連続的・集中的・選択的な対応によって成し遂げられる。6） クロマチン構造の可塑性の不可逆的創出は、体細胞のそれまでの履歴と環境変化への連続的対応に依存する。7） クロマチン構造の可塑性の不可逆的創出は、子孫世代の細胞で起り、環境変化に最初に遭遇した細胞では起らない。8） クロマチン構造の可塑性の不可逆的創出は、合目的的・必然的に起り、偶然または中立的に起るのではない。9） 環境変化は環境変化認識受容体/部位（environment change recognition receptor/site: ECRR/ECRS）によって認識される。10） クロマチン構造の可塑性はクロマチン・コンフォメーション・チェンジ・コンプレックス装置（chromatin conformation change complex machinery: 4C machinery）によって直接的・不可逆的に創出される。11） 特定の遺伝子（群）の5'-近傍上流のクロマチン構造は、動的で変化可能な三次元構造（コンフォメーション）として、環境変化のシグナルを受け取る。12） 特定の遺伝子の5'-近傍上流のクロマチン構造（緩いフォームまたは硬いフォーム）は当該遺伝子の潜在的な発現能力のスイッチ（オンまたはオフ）を指令する。13） 特定の遺伝子の5'-近傍上流のクロマチン構造は、構造的側面からは「クロマチンの刻み目（notch of chromatin）」とみなすことができ、機能的側面からは「遺伝子発現のディレクター（director of gene expression）」とみなすことができる。14） 高等真核生物の複雑で多彩な細胞機能や多種多様な細胞タイプを決定する4C theory でのコード数は、恐らく特異的な候補遺伝子数とこれら遺伝子それぞれのコード数（多分、2）の組合せに基づいて決まる。","subitem_description_language":"ja","subitem_description_type":"Abstract"}]},"item_10010_version_type_20":{"attribute_name":"著者版フラグ","attribute_value_mlt":[{"subitem_version_resource":"http://purl.org/coar/version/c_ab4af688f83e57aa","subitem_version_type":"AM"}]},"item_creator":{"attribute_name":"著者","attribute_type":"creator","attribute_value_mlt":[{"creatorNames":[{"creatorName":"中山, 建男","creatorNameLang":"ja"},{"creatorName":"ナカヤマ, タツオ","creatorNameLang":"ja-Kana"},{"creatorName":"Nakayama, Tatsuo","creatorNameLang":"en"}],"nameIdentifiers":[{}]},{"creatorNames":[{"creatorName":"中山, 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