£€0€ €0€ €0€ €­t¢€ €¡€0€ €4JSX£€0€¢€0€ €Protein Data Bank¡€ €¡€0€ €è¡€¢€¢€¡€0€ €PDB¡€¡€4JSX£€¡€0€ €Ý¡€¢€¤€¡€0€ €0€ € €0€0€ €¡€0€ € Pavletich£€ N.P.Pavletich¤€N.P.0€ €¡€0€ €Yang£€H.Yang¤€H.¡€0€ €¡€0€ €Ý¡€¢€¤€«€1€¤€0€ €1€ €0mTOR kinase structure, mechanism and regulation.¡€0€ € €0€0€ €¡€0€ €Yang£€H.Yang¤€H.0€ €¡€0€ €Rudge£€ D.G.Rudge¤€D.G.0€ €¡€0€ €Koos£€J.D.Koos¤€J.D.0€ €¡€0€ € Vaidialingam£€B.Vaidialingam¤€B.0€ €¡€0€ €Yang£€H.J.Yang¤€H.J.0€ €¡€0€ € Pavletich£€ N.P.Pavletich¤€N.P.¢€ €0€ €1€¤€Nature¥€Nature €Nature¡€0€ €¡€0€ €Ý¡€¢€ ¡€497¢€7448£€217-23¨€eng£€1€ €h©f¢€10.1038/nature12122£€ nature12122¥€ PMC4512754§€0€ €PDB Citation Status¡€¡€primary¢€0€¡€¡€0€ €Ý¡€¢€ ¢€0€¢€ €0€ €1€ €0mTOR kinase structure, mechanism and regulation.¡€0€ €¡€0€ €Ý¡€¢€ £€‚The mammalian target of rapamycin (mTOR), a phosphoinositide 3-kinase-related protein kinase, controls cell growth in response to nutrients and growth factors and is frequently deregulated in cancer. Here we report co-crystal structures of a complex of truncated mTOR and mammalian lethal with SEC13 protein 8 (mLST8) with an ATP transition state mimic and with ATP-site inhibitors. The structures reveal an intrinsically active kinase conformation, with catalytic residues and a catalytic mechanism remarkably similar to canonical protein kinases. The active site is highly recessed owing to the FKBP12-rapamycin-binding (FRB) domain and an inhibitory helix protruding from the catalytic cleft. mTOR-activating mutations map to the structural framework that holds these elements in place, indicating that the kinase is controlled by restricted access. In vitro biochemistry shows that the FRB domain acts as a gatekeeper, with its rapamycin-binding site interacting with substrates to grant them access to the restricted active site. Rapamycin-FKBP12 inhibits the kinase by directly blocking substrate recruitment and by further restricting active-site access. The structures also reveal active-site residues and conformational changes that underlie inhibitor potency and specificity.¬€h©f¢€0€ €0€ €,structure of mTORDeltaN-mLST8-Torin2 complex¡€ TRANSFERASE¢€‚Mol_id: 1 Organism_scientific: Homo sapiens Organism_taxid: 9606 Mol_id: 2 Organism_scientific: Homo sapiens Organism_taxid: 9606£€‚ƒstructure of mTORDeltaN-mLST8-Torin2 complex; kinase, transferase; Mol_id: 1; Molecule: Serine/threonine-protein kinase mTOR; Chain: B, A; Fragment: unp residues 1376-2549; Synonym: FK506-binding protein 12-rapamycin complex-associated protein 1, FKBP12-rapamycin complex-associated protein, Mammalian target of rapamycin, mTOR, Mechanistic target of rapamycin, Rapamycin and FKBP12 target 1, Rapamycin target protein 1; EC number: 2.7.11.1; Mol_id: 2; Molecule: Target of rapamycin complex subunit LST8; Chain: D, C; Synonym: TORC subunit LST8, G protein beta subunit-like, Gable, Protein GbetaL, Mammalian lethal with SEC13 protein 8, mLST8;¡€0€0€ €¡€0€ €B£€&1 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