+データを開く
-基本情報
登録情報 | データベース: PDB / ID: 8ppl | ||||||||||||||||||||||||||||||||||||||||||
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タイトル | MERS-CoV Nsp1 bound to the human 43S pre-initiation complex | ||||||||||||||||||||||||||||||||||||||||||
要素 |
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キーワード | TRANSLATION / Nsp1 / MERS / SARS / SARS-CoV2 / ribosome / 40S ribosomal subunit / translation inhibition / coronavirus / 43S PIC / 43S pre-initiation complex / mRNA channel / initiation factor / eIF2 / eIF3 / eIF1 / eIF1A / VIRAL PROTEIN | ||||||||||||||||||||||||||||||||||||||||||
機能・相同性 | 機能・相同性情報 male germ cell proliferation / positive regulation of mRNA binding / regulation of translation in response to endoplasmic reticulum stress / translation initiation ternary complex / positive regulation of mRNA cis splicing, via spliceosome / glial limiting end-foot / response to kainic acid / HRI-mediated signaling / Cellular response to mitochondrial stress / viral translational termination-reinitiation ...male germ cell proliferation / positive regulation of mRNA binding / regulation of translation in response to endoplasmic reticulum stress / translation initiation ternary complex / positive regulation of mRNA cis splicing, via spliceosome / glial limiting end-foot / response to kainic acid / HRI-mediated signaling / Cellular response to mitochondrial stress / viral translational termination-reinitiation / response to manganese-induced endoplasmic reticulum stress / positive regulation of type B pancreatic cell apoptotic process / methionyl-initiator methionine tRNA binding / eukaryotic translation initiation factor 3 complex, eIF3e / negative regulation of translational initiation in response to stress / Response of EIF2AK1 (HRI) to heme deficiency / cap-dependent translational initiation / eukaryotic translation initiation factor 3 complex, eIF3m / Recycling of eIF2:GDP / PERK-mediated unfolded protein response / translation reinitiation / PERK regulates gene expression / eukaryotic translation initiation factor 2 complex / IRES-dependent viral translational initiation / regulation of translational initiation in response to stress / multi-eIF complex / formation of cytoplasmic translation initiation complex / eukaryotic translation initiation factor 3 complex / protein-synthesizing GTPase / eukaryotic 43S preinitiation complex / cytoplasmic translational initiation / translation factor activity, RNA binding / mRNA cap binding / formation of translation preinitiation complex / positive regulation of cysteine-type endopeptidase activity involved in execution phase of apoptosis / negative regulation of endoplasmic reticulum unfolded protein response / oxidized pyrimidine DNA binding / response to TNF agonist / positive regulation of base-excision repair / eukaryotic 48S preinitiation complex / protein tyrosine kinase inhibitor activity / positive regulation of respiratory burst involved in inflammatory response / regulation of adenylate cyclase-activating G protein-coupled receptor signaling pathway / positive regulation of intrinsic apoptotic signaling pathway in response to DNA damage / positive regulation of gastrulation / nucleolus organization / IRE1-RACK1-PP2A complex / positive regulation of endodeoxyribonuclease activity / positive regulation of Golgi to plasma membrane protein transport / TNFR1-mediated ceramide production / negative regulation of DNA repair / negative regulation of RNA splicing / metal-dependent deubiquitinase activity / negative regulation of intrinsic apoptotic signaling pathway in response to hydrogen peroxide / oxidized purine DNA binding / supercoiled DNA binding / neural crest cell differentiation / NF-kappaB complex / ubiquitin-like protein conjugating enzyme binding / regulation of translational initiation / regulation of establishment of cell polarity / negative regulation of phagocytosis / positive regulation of ubiquitin-protein transferase activity / rRNA modification in the nucleus and cytosol / Formation of the ternary complex, and subsequently, the 43S complex / erythrocyte homeostasis / cytoplasmic side of rough endoplasmic reticulum membrane / laminin receptor activity / nuclear-transcribed mRNA catabolic process, nonsense-mediated decay / protein kinase A binding / negative regulation of ubiquitin protein ligase activity / Ribosomal scanning and start codon recognition / ion channel inhibitor activity / Translation initiation complex formation / pigmentation / positive regulation of mitochondrial depolarization / mammalian oogenesis stage / activation-induced cell death of T cells / negative regulation of Wnt signaling pathway / positive regulation of T cell receptor signaling pathway / fibroblast growth factor binding / positive regulation of activated T cell proliferation / iron-sulfur cluster binding / regulation of cell division / Protein hydroxylation / monocyte chemotaxis / negative regulation of peptidyl-serine phosphorylation / BH3 domain binding / mTORC1-mediated signalling / SARS-CoV-1 modulates host translation machinery / Peptide chain elongation / positive regulation of intrinsic apoptotic signaling pathway by p53 class mediator / cysteine-type endopeptidase activator activity involved in apoptotic process / Selenocysteine synthesis / positive regulation of signal transduction by p53 class mediator / Formation of a pool of free 40S subunits / Eukaryotic Translation Termination / phagocytic cup / ubiquitin ligase inhibitor activity / Response of EIF2AK4 (GCN2) to amino acid deficiency 類似検索 - 分子機能 | ||||||||||||||||||||||||||||||||||||||||||
生物種 | Middle East respiratory syndrome-related coronavirus (ウイルス) Homo sapiens (ヒト) | ||||||||||||||||||||||||||||||||||||||||||
手法 | 電子顕微鏡法 / 単粒子再構成法 / クライオ電子顕微鏡法 / 解像度: 2.65 Å | ||||||||||||||||||||||||||||||||||||||||||
データ登録者 | Schubert, K. / Karousis, E.D. / Ban, I. / Lapointe, C.P. / Leibundgut, M. / Baeumlin, E. / Kummerant, E. / Scaiola, A. / Schoenhut, T. / Ziegelmueller, J. ...Schubert, K. / Karousis, E.D. / Ban, I. / Lapointe, C.P. / Leibundgut, M. / Baeumlin, E. / Kummerant, E. / Scaiola, A. / Schoenhut, T. / Ziegelmueller, J. / Puglisi, J.D. / Muehlemann, O. / Ban, N. | ||||||||||||||||||||||||||||||||||||||||||
資金援助 | スイス, 米国, 13件
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引用 | ジャーナル: Mol Cell / 年: 2023 タイトル: Universal features of Nsp1-mediated translational shutdown by coronaviruses. 著者: Katharina Schubert / Evangelos D Karousis / Ivo Ban / Christopher P Lapointe / Marc Leibundgut / Emilie Bäumlin / Eric Kummerant / Alain Scaiola / Tanja Schönhut / Jana Ziegelmüller / ...著者: Katharina Schubert / Evangelos D Karousis / Ivo Ban / Christopher P Lapointe / Marc Leibundgut / Emilie Bäumlin / Eric Kummerant / Alain Scaiola / Tanja Schönhut / Jana Ziegelmüller / Joseph D Puglisi / Oliver Mühlemann / Nenad Ban / 要旨: Nonstructural protein 1 (Nsp1) produced by coronaviruses inhibits host protein synthesis. The severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) Nsp1 C-terminal domain was shown to bind the ...Nonstructural protein 1 (Nsp1) produced by coronaviruses inhibits host protein synthesis. The severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) Nsp1 C-terminal domain was shown to bind the ribosomal mRNA channel to inhibit translation, but it is unclear whether this mechanism is broadly used by coronaviruses, whether the Nsp1 N-terminal domain binds the ribosome, or how Nsp1 allows viral RNAs to be translated. Here, we investigated Nsp1 from SARS-CoV-2, Middle East respiratory syndrome coronavirus (MERS-CoV), and Bat-Hp-CoV coronaviruses using structural, biophysical, and biochemical experiments, revealing a conserved role for the C-terminal domain. Additionally, the N-terminal domain of Bat-Hp-CoV Nsp1 binds to the decoding center of the 40S subunit, where it would prevent mRNA and eIF1A accommodation. Structure-based experiments demonstrated the importance of decoding center interactions in all three coronaviruses and showed that the same regions of Nsp1 are necessary for the selective translation of viral RNAs. Our results provide a mechanistic framework to understand how Nsp1 controls preferential translation of viral RNAs. #1: ジャーナル: Mol.Cell / 年: 2023 タイトル: Universal features of Nsp1-mediated translational shutdown by coronaviruses 著者: Schubert, K. / Karousis, E.D. / Ban, I. / Lapointe, C.P. / Leibundgut, M. / Baeumlin, E. / Kummerant, E. / Scaiola, A. / Schoenhut, T. / Ziegelmueller, J. / Puglisi, J.D. / Muehlemann, O. / Ban, N. | ||||||||||||||||||||||||||||||||||||||||||
履歴 |
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-構造の表示
構造ビューア | 分子: MolmilJmol/JSmol |
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-ダウンロードとリンク
-ダウンロード
PDBx/mmCIF形式 | 8ppl.cif.gz | 3 MB | 表示 | PDBx/mmCIF形式 |
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PDB形式 | pdb8ppl.ent.gz | 表示 | PDB形式 | |
PDBx/mmJSON形式 | 8ppl.json.gz | ツリー表示 | PDBx/mmJSON形式 | |
その他 | その他のダウンロード |
-検証レポート
文書・要旨 | 8ppl_validation.pdf.gz | 1.9 MB | 表示 | wwPDB検証レポート |
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文書・詳細版 | 8ppl_full_validation.pdf.gz | 2 MB | 表示 | |
XML形式データ | 8ppl_validation.xml.gz | 256 KB | 表示 | |
CIF形式データ | 8ppl_validation.cif.gz | 418.1 KB | 表示 | |
アーカイブディレクトリ | https://data.pdbj.org/pub/pdb/validation_reports/pp/8ppl ftp://data.pdbj.org/pub/pdb/validation_reports/pp/8ppl | HTTPS FTP |
-関連構造データ
関連構造データ | 17805MC 8ppkC C: 同じ文献を引用 (文献) M: このデータのモデリングに利用したマップデータ |
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類似構造データ | 類似検索 - 機能・相同性F&H 検索 |
-リンク
-集合体
登録構造単位 |
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1 |
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-要素
-Eukaryotic translation initiation factor ... , 15種, 15分子 I3I4I5I6I8IoIpIqIrIsItIuIvIxIy
#1: タンパク質 | 分子量: 25083.619 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: Q9UBQ5 |
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#2: タンパク質 | 分子量: 37593.645 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: O00303, ubiquitinyl hydrolase 1 |
#3: タンパク質 | 分子量: 66803.734 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: Q9Y262 |
#4: タンパク質 | 分子量: 42555.832 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: Q7L2H7 |
#5: タンパク質 | 分子量: 39979.277 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: O15372 |
#6: タンパク質 | 分子量: 35662.016 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: O75821 |
#7: タンパク質 | 分子量: 12752.498 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: N-terminus built as UNK / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P41567 |
#8: タンパク質 | 分子量: 16488.449 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P47813 |
#9: タンパク質 | 分子量: 36161.180 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P05198 |
#10: タンパク質 | 分子量: 38454.484 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P20042 |
#11: タンパク質 | 分子量: 51178.406 Da / 分子数: 1 / 由来タイプ: 天然 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P41091, protein-synthesizing GTPase |
#12: タンパク質 | 分子量: 166903.781 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: UniProt accession code Q14152 / 由来: (天然) Homo sapiens (ヒト) / 参照: UniProt: Q14152 |
#13: タンパク質 | 分子量: 52281.633 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: UniProt accession code P60228 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P60228 |
#15: タンパク質 | 分子量: 64060.758 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: UniProt accession code O15371 / 由来: (天然) Homo sapiens (ヒト) / 参照: UniProt: O15371 |
#16: タンパク質 | 分子量: 105503.945 Da / 分子数: 1 / 由来タイプ: 天然 詳細: One linker built as UNK. UniProt accession code Q99613 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: Q99613 |
-RNA鎖 , 2種, 2分子 IwA2
#14: RNA鎖 | 分子量: 24463.807 Da / 分子数: 1 / 由来タイプ: 天然 詳細: Aminoacylated initiator tRNA with base modifications. (XXX) refers to methionine residue attached to CCA-3'OH of tRNA. Sequence obtained from gtrnadb. 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E |
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#17: RNA鎖 | 分子量: 603609.188 Da / 分子数: 1 / 由来タイプ: 天然 詳細: unmodified sequence: >18S_rRNA_taoka_unmodified UACCUGGUUGAUCCUGCCAGUAGCAUaUGCUUGuCuCAAAGAUUAAGCCAUGCAUGUCUA AGUACGCACGGCCGGUACAGUGAAACUGCGAAuGGCUCaUUAAAuCAGuUAUGGUuCCuU ...詳細: unmodified sequence: >18S_rRNA_taoka_unmodified UACCUGGUUGAUCCUGCCAGUAGCAUaUGCUUGuCuCAAAGAUUAAGCCAUGCAUGUCUA AGUACGCACGGCCGGUACAGUGAAACUGCGAAuGGCUCaUUAAAuCAGuUAUGGUuCCuU uGGUCGCUCGCUCCUCUCCUACUUGGAUAACUGUGGUAaUUCUAGaGCUAAuAcAUGCCG ACGGGCGCUGACCCCCUUCGCGGGGGGGAuGCGUGCAuUUAUCAGAUCAAAACCAACCCG GUCAGCCCCUCUCCGGCCCCGGCCGGGGGGCGGGCGCCGGCGGCUUUGGUGACUCuAGAU AACCUCGGGCCGAUCGCACGCCCCCCGUGGCGGCGACGACCCAUUCGAACGUCuGCCCUA UCAACUUUCGAUGGUAGUCGCCGUGCCUACCAUGGUGACCACGGGuGACGGGGAAUCAGG GUUCGAUuCCGGAGAgGGAGCCUGAGAAACGGCUACCACAUcCAAGGaAGGCAGCAGGCG CGCaAAUUACCCACUCCCGACCCGGGGAgGUaGUGAcGAAAAAUAACAAUACAGGACUCU UUCGAGGCCCUGUAAUUGGAAUGAGUCCACUuUAAaUCCUUUAACGAGGaUCCAUUGGAG gGCAAGUCuGGUGCCAGCAGcCGCGGuAAUUCCAGCUCCAAUAgCGUAuAuUAAAGUUGC UGCAGUUaAAAAGCUCGUAGuUgGAuCUUGGGAGCGGGCGGGCGGUCCGCCGCGAGGCGA GCCACCGCCCGUCCCCGCCCCUUGCCUCUCGGCGCCCCCUCGAUGCUCUUAGCUGAGUGU CCCGCGGGGCCCGAAGcGuUuACUUUGAAAAAAuuAGAGUGuUCAAAGCAGGCCCGAGCC GCCUGGAUACCGCAGCUAGGAAuAAugGAAUAGGACCGCGGUUCUAUUUUGUUGGUuUUC GGAACUGAGGCCAUGAUuAAGAGGGACGGCCGGGGGCAUUCGUAUUGCGCCGCUAGAGGU GAAAUuCUUGGACCGGCGCAAGACGGACCAGAGCGAAAGCAUUuGCCAAGAAUGUUUUCA UUAAUCAAGAaCGAAAGUCGGAGGuuCGAAGACGAuCAGAUACCGUCGUAGUUCCGACCA uAAACGAUGCCGACCGGCGAUGCGGCGGCGUUAUUCCCAUGACCCGCCGGGCAGCuUCCG GGAAACCAAAGUCUUUGGGUUCCGGGGGGAGUAuGGuUGCAAAGCUGAAACUUAAAGGAA UUGACGGAAGGGCACCACCAGGAGUGGAGCCuGCGGCuUAAUUuGACuCAACACGGGAAA CCUCACCCGGCcCGGACACGGACAGGAuUGACAGAUUGAUAGCUCUUUCUCGAUUCCGUG GGUGGuGgUGCAUGGCcGUUCUUAGUuGGUGGAGCGAUUUGUCUGGuUAAUUCCGAUAAC GAaCGAGACUcUGGCAUGCUAACUAGUUACGCGACCCCCGAGCGGUCGGCGUCCCCCAAC UuCUuAgAGGGACAAGUGGCGUUCAGCCACCCGAGAUUGAGCAAUAACAgGUCUGUGAUG CCCUUAGAUGUCCGGGGCUGCACGCGCGCUACACUGACUGGCUCAGCGUGUGCCUACCCU ACGCCGGCAGGCGCGGGUAACCCGUUGAACCCCAUUCGUGAUGGGGAUCGGGGAUUGCAA UUAUuCCCCAUGAACGAGgAAUuCCCAGUAAGUGCGGGUCAUAAGCUuGCGUUGAUUaAG UCCCUGCCCUUuGUACACACCGcCCGUCGCUACUACCGAUUGGAUGGUUUAGUGAGGCCC UCGGAUCGGCCCCGCCGGGGUCGGCCCACGGCCCUGGCGGAGCGCUGAGAAGACGGUCGA ACUuGACUAUCUAGAGGAAGUAAAAGUCGUAaCAAGGUUUCcGUAGGUGaaCCUGCGGAA GGAUCAUUA 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E |
+40S ribosomal protein ... , 28種, 28分子 AAABACADAEAFAGAHAIAJAKALAMANAOAPAQARAUAVAWAXAYAZAaAbAcAd
-Small ribosomal subunit protein ... , 2種, 2分子 ASAT
#36: タンパク質 | 分子量: 17801.814 Da / 分子数: 1 / 由来タイプ: 天然 詳細: UniProt accession code P62269 S2: acetylserine (SAC) 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P62269 |
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#37: タンパク質 | 分子量: 16104.579 Da / 分子数: 1 / 由来タイプ: 天然 詳細: R67: omega-methylarginine (NMM) UniProt accession code P39019 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P39019 |
-タンパク質 , 4種, 4分子 AeAfAgAj
#48: タンパク質 | 分子量: 14415.724 Da / 分子数: 1 / 由来タイプ: 天然 詳細: P62861 (UniProt) NP_001988.1 (GeneBank) residues 1-74: ubiquitin 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P62861 |
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#49: タンパク質 | 分子量: 18004.041 Da / 分子数: 1 / 由来タイプ: 天然 詳細: residues 1-76: ubiquitin zinc finger protein UniProt accession code P62979 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P62979 |
#50: タンパク質 | 分子量: 35115.652 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: UniProt accession code P63244 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293E / 参照: UniProt: P63244 |
#52: タンパク質 | 分子量: 24328.531 Da / 分子数: 1 / 由来タイプ: 組換発現 詳細: residues 1-193 of MERS polyprotein ORF1ab UniProt accession number K9N7C7 (R1AB_MERS1) 由来: (組換発現) Middle East respiratory syndrome-related coronavirus (ウイルス) 遺伝子: rep, 1a-1b / プラスミド: pcDNA3.1(+) / 細胞株 (発現宿主): HEK293E / 発現宿主: Homo sapiens (ヒト) / 参照: UniProt: K9N7C7 |
-タンパク質・ペプチド , 1種, 1分子 Ah
#51: タンパク質・ペプチド | 分子量: 3473.451 Da / 分子数: 1 / 由来タイプ: 天然 / 詳細: UniProt accession code P62945 / 由来: (天然) Homo sapiens (ヒト) / 細胞株: HEK293 / 参照: UniProt: P62945 |
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-非ポリマー , 6種, 253分子
#53: 化合物 | ChemComp-ZN / #54: 化合物 | ChemComp-GTP / | #55: 化合物 | ChemComp-MG / #56: 化合物 | ChemComp-MET / | #57: 化合物 | ChemComp-UNX / #58: 水 | ChemComp-HOH / | |
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-詳細
研究の焦点であるリガンドがあるか | N |
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-実験情報
-実験
実験 | 手法: 電子顕微鏡法 |
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EM実験 | 試料の集合状態: PARTICLE / 3次元再構成法: 単粒子再構成法 |
-試料調製
構成要素 | 名称: MERS-CoV Nsp1 - 43S pre-initiation complex / タイプ: RIBOSOME / Entity ID: #1-#13, #15-#52 / 由来: NATURAL | ||||||||||||||||
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由来(天然) | 生物種: Homo sapiens (ヒト) | ||||||||||||||||
緩衝液 | pH: 7.4 | ||||||||||||||||
緩衝液成分 |
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試料 | 包埋: NO / シャドウイング: NO / 染色: NO / 凍結: YES / 詳細: f.c. 100 nM | ||||||||||||||||
試料支持 | 詳細: 15 sec in easiGlow Discharge cleaning system (PELCO) at 15 mA グリッドの材料: COPPER / グリッドのタイプ: Quantifoil R2/2 | ||||||||||||||||
急速凍結 | 装置: FEI VITROBOT MARK IV / 凍結剤: ETHANE-PROPANE / 湿度: 100 % / 凍結前の試料温度: 277 K |
-電子顕微鏡撮影
実験機器 | モデル: Titan Krios / 画像提供: FEI Company |
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顕微鏡 | モデル: FEI TITAN KRIOS |
電子銃 | 電子線源: FIELD EMISSION GUN / 加速電圧: 300 kV / 照射モード: FLOOD BEAM |
電子レンズ | モード: BRIGHT FIELD / 倍率(公称値): 81000 X / 最大 デフォーカス(公称値): 3000 nm / 最小 デフォーカス(公称値): 600 nm |
試料ホルダ | 凍結剤: NITROGEN 試料ホルダーモデル: FEI TITAN KRIOS AUTOGRID HOLDER |
撮影 | 電子線照射量: 60 e/Å2 / フィルム・検出器のモデル: GATAN K3 (6k x 4k) / 実像数: 8932 |
-解析
EMソフトウェア |
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CTF補正 | タイプ: PHASE FLIPPING AND AMPLITUDE CORRECTION | |||||||||||||||
3次元再構成 | 解像度: 2.65 Å / 解像度の算出法: FSC 0.143 CUT-OFF / 粒子像の数: 218516 / 対称性のタイプ: POINT | |||||||||||||||
原子モデル構築 | プロトコル: OTHER / 空間: REAL / 詳細: phenix.real_space_refine | |||||||||||||||
原子モデル構築 | 3D fitting-ID: 1 / Source name: PDB / タイプ: experimental model
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