5ZCS: Serine/threonine-protein kinase mTOR

4.9 Angstrom Cryo-EM structure of human mTOR complex 2. Determined by electron microscopy at 4.9 Å resolution. Released 21 Mar 2018.

Method: Electron microscopy
Resolution: 4.9 Å
Organism: Homo sapiens
Chains: 8
Atoms: 49,176
Mol. weight: 1110.94 kDa
Released: 21 Mar 2018

Explore 5ZCS in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

5ZCS contains 420 α-helices and 86 β-strands across 8 chains. Residue ranges use author residue numbering, as in the PDB file, from PDBe. To see them in 3D, choose the Cartoon representation with Secondary structure coloring: helices, sheets and coils get different colors.

Chains A and B: 154 helices, 10 β-strands

Element	Residues	Length	Sheet
α-helix	26-29	4
α-helix	64-73	10
α-helix	86-93	8
α-helix	102-107	6
α-helix	108-110	3
α-helix	122-131	10
α-helix	141-150	10
α-helix	166-171	6
α-helix	181-183	3
α-helix	190-198	9
α-helix	210-214	5
α-helix	218-222	5
α-helix	235-245	11
α-helix	267-273	7
α-helix	277-284	8
α-helix	357-360	4
α-helix	363-374	12
α-helix	389-392	4
α-helix	396-402	7
α-helix	413-420	8
α-helix	431-437	7
α-helix	448-450	3
α-helix	452-457	6
α-helix	482-487	6
α-helix	499-503	5
α-helix	505-508	4
α-helix	518-525	8
α-helix	532-542	11
α-helix	544-547	4
α-helix	580-589	10
α-helix	604-606	3
α-helix	622-629	8
α-helix	649-663	15
α-helix	669-674	6
α-helix	681-683	3
α-helix	690-697	8
α-helix	711-717	7
α-helix	729-732	4
α-helix	747-756	10
α-helix	765-767	3
α-helix	772-777	6
α-helix	781-783	3
α-helix	792-802	11
α-helix	814-816	3
α-helix	825-828	4
α-helix	837-846	10
α-helix	848-851	4
α-helix	867-874	8
α-helix	878-890	13
α-helix	896-902	7
α-helix	937-939	3
α-helix	943-946	4
α-helix	969-980	12
α-helix	994-997	4
α-helix	1000-1003	4
α-helix	1012-1019	8
α-helix	1028-1030	3
α-helix	1032-1037	6
α-helix	1046-1052	7
α-helix	1060-1063	4
α-helix	1067-1069	3
α-helix	1072-1074	3
α-helix	1075-1079	5
α-helix	1091-1100	10
α-helix	1120-1122	3
α-helix	1138-1143	6
α-helix	1153-1155	3
α-helix	1158-1165	8
α-helix	1167-1169	3
α-helix	1170-1173	4
α-helix	1176-1184	9
α-helix	1189-1191	3
α-helix	1192-1202	11
α-helix	1209-1218	10
α-helix	1263-1266	4
α-helix	1277-1293	17
α-helix	1298-1300	3
α-helix	1304-1307	4
α-helix	1311-1315	5
α-helix	1320-1326	7
α-helix	1334-1342	9
α-helix	1351-1359	9
α-helix	1374-1376	3
α-helix	1382-1389	8
α-helix	1400-1404	5
α-helix	1426-1429	4
α-helix	1448-1452	5
α-helix	1460-1467	8
α-helix	1474-1480	7
α-helix	1482-1485	4
α-helix	1490-1497	8
α-helix	1506-1521	16
α-helix	1527-1533	7
α-helix	1545-1553	9
α-helix	1557-1565	9
α-helix	1566-1569	4
α-helix	1573-1576	4
α-helix	1587-1596	10
α-helix	1612-1618	7
α-helix	1623-1625	3
α-helix	1630-1632	3
α-helix	1634-1636	3
α-helix	1639-1641	3
α-helix	1646-1648	3
α-helix	1653-1661	9
α-helix	1666-1677	12
α-helix	1694-1706	13
α-helix	1713-1724	12
α-helix	1727-1730	4
α-helix	1737-1750	14
α-helix	1751-1754	4
α-helix	1758-1761	4
α-helix	1769-1780	12
α-helix	1787-1789	3
α-helix	1792-1807	16
α-helix	1808-1812	5
α-helix	1870-1877	8
α-helix	1886-1892	7
α-helix	1899-1902	4
α-helix	1905-1913	9
α-helix	1917-1922	6
α-helix	1924-1927	4
α-helix	1957-1964	8
α-helix	1969-1971	3
α-helix	1973-1976	4
α-helix	1985-1997	13
α-helix	1998-2000	3
α-helix	2006-2020	15
α-helix	2023-2040	18
α-helix	2046-2055	10
α-helix	2065-2073	9
α-helix	2078-2086	9
α-helix	2098-2112	15
α-helix	2123-2126	4
α-helix	2128-2130	3
β-strand	2139	1	1
β-strand	2152	1	1
β-strand	2156	1	2
β-strand	2170-2175	6	2
β-strand	2181-2187	7	2
α-helix	2201-2208	8
α-helix	2213-2216	4
α-helix	2227	1
β-strand	2228-2229	2	2
β-strand	2235-2237	3	2
α-helix	2239-2240	2
β-strand	2243-2244	2	3
α-helix	2247-2253	7
α-helix	2266-2271	6
α-helix	2288-2290	3
α-helix	2299-2302	4
α-helix	2311-2322	12
α-helix	2330-2334	5
α-helix	2341-2343	3
β-strand	2344-2347	4	3
β-strand	2353-2355	3	3
α-helix	2381-2385	5
α-helix	2397-2408	12
α-helix	2411-2414	4
α-helix	2418-2421	4
α-helix	2428-2430	3
α-helix	2493-2509	17
α-helix	2521-2527	7
α-helix	2530-2533	4

Chains C and D: 0 helices, 33 β-strands

Element	Residues	Length	Sheet
β-strand	13-18	6	7
β-strand	23	1	8
β-strand	24-27	4	7
β-strand	39	1	8
β-strand	45-50	6	9
β-strand	56-61	6	9
β-strand	67-69	3	9
β-strand	90-93	4	10
β-strand	99-103	5	10
β-strand	109	1	10
β-strand	112-113	2	10
β-strand	131-136	6	11
β-strand	142-147	6	11
β-strand	152-156	5	11
β-strand	163-165	3	11
β-strand	178	1	12
β-strand	184-188	5	12
β-strand	193-198	6	12
β-strand	211-217	7	12
β-strand	223-228	6	13
β-strand	234-239	6	13
β-strand	243-245	3	13
β-strand	247-248	2	13
β-strand	257-259	3	13
β-strand	275	1	14
β-strand	278	1	15
β-strand	284	1	16
β-strand	285	1	15
β-strand	288	1	14
β-strand	294-295	2	17
β-strand	298	1	16
β-strand	307-308	2	17
β-strand	318-322	5	7

Chains E and F: 52 helices, 0 β-strands

Element	Residues	Length
α-helix	193-210	18
α-helix	217-233	17
α-helix	241-249	9
α-helix	258-267	10
α-helix	274-277	4
α-helix	283-291	9
α-helix	293-296	4
α-helix	299-315	17
α-helix	321-326	6
α-helix	328-336	9
α-helix	344-353	10
α-helix	366-373	8
α-helix	379-398	20
α-helix	406-408	3
α-helix	411-417	7
α-helix	421-423	3
α-helix	431-447	17
α-helix	457-468	12
α-helix	474-477	4
α-helix	478-482	5
α-helix	499-506	8
α-helix	514-535	22
α-helix	542-548	7
α-helix	552-556	5
α-helix	565-577	13
α-helix	582-591	10
α-helix	595-608	14
α-helix	610-612	3
α-helix	623-631	9
α-helix	637-639	3
α-helix	653-664	12
α-helix	676-680	5
α-helix	694-704	11
α-helix	732-743	12
α-helix	752-763	12
α-helix	765-771	7
α-helix	802-810	9
α-helix	817-823	7
α-helix	830-842	13
α-helix	848-860	13
α-helix	868-871	4
α-helix	881-884	4
α-helix	891-895	5
α-helix	897-906	10
α-helix	912-929	18
α-helix	938-957	20
α-helix	960-974	15
α-helix	986-993	8
α-helix	1003-1017	15
α-helix	1611-1625	15
α-helix	1631-1647	17
α-helix	1681-1694	14

Chains G and H: 4 helices, 0 β-strands

Element	Residues	Length
α-helix	10-19	10
α-helix	50-57	8
α-helix	66-75	10
α-helix	91-103	13

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Serine/threonine-protein kinase mTOR	A, B	protein	2549	Homo sapiens	P42345 (AlphaFold model)
Target of rapamycin complex subunit LST8	C, D	protein	326	Homo sapiens	Q9BVC4 (AlphaFold model)
Rapamycin-insensitive companion of mTOR	E, F	protein	1708	Homo sapiens	Q6R327 (AlphaFold model)
Target of rapamycin complex 2 subunit MAPKAP1	G, H	protein	522	Homo sapiens	Q9BPZ7 (AlphaFold model)

Sequence of entity 1 (A, B), FASTA

>5ZCS_1 Serine/threonine-protein kinase mTOR (chains A, B)
MLGTGPAAATTAATTSSNVSVLQQFASGLKSRNEETRAKAAKELQHYVTMELREMSQEES
TRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDP
VVMEMASKAIGRLAMAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVP
TFFFQQVQPFFDNIFVAVWDPKQAIREGAVAALRACLILTTQREPKEMQKPQWYRHTFEE
AEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQLVHDKYC
KDLMGFGTKPRHITPFTSFQAVQPQQSNALVGLLGYSSHQGLMGFGTSPSPAKSTLVESR
CCRDLMEEKFDQVCQWVLKCRNSKNSLIQMTILNLLPRLAAFRPSAFTDTQYLQDTMNHV
LSCVKKEKERTAAFQALGLLSVAVRSEFKVYLPRVLDIIRAALPPKDFAHKRQKAMQVDA
TVFTCISMLARAMGPGIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLL
KMLSLVLMHKPLRHPGMPKGLAHQLASPGLTTLPEASDVGSITLALRTLGSFEFEGHSLT
QFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLL
VVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSS
MNPAFVMPFLRKMLIQILTELEHSGIGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALI
LKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVA
LWTLGQLVASTGYVVEPYRKYPTLLEVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHK
VNIGMIDQSRDASAVSLSESKSSQDSSDYSTSEMLVNMGNLPLDEFYPAVSMVALMRIFR
DQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLV
SFVKSHIRPYMDEIVTLMREFWVMNTSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLR
VFMHDNSPGRIVSIKLLAAIQLFGANLDDYLHLLLPPIVKLFDAPEAPLPSRKAALETVD
RLTESLDFTDYASRIIHPIVRTLDQSPELRSTAMDTLSSLVFQLGKKYQIFIPMVNKVLV
RHRINHQRYDVLICRIVKGYTLADEEEDPLIYQHRMLRSGQGDALASGPVETGPMKKLHV
STINLQKAWGAARRVSKDDWLEWLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNA
AFVSCWSELNEDQQDELIRSIELALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGI
VLLGERAAKCRAYAKALHYKELEFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHF
GELEIQATWYEKLHEWEDALVAYDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEK
WTLVNDETQAKMARMAAAAAWGLGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLA
QQCIDKARDLLDAELTAMAGESYSRAYGAMVSCHMLSELEEVIQYKLVPERREIIRQIWW
ERLQGCQRIVEDWQKILMVRSLVVSPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVD
PSRQLDHPLPTVHPQVTYAYMKNMWKSARKIDAFQHMQHFVQTMQQQAQHAIATEDQQHK
QELHKLMARCFLKLGEWQLNLQGINESTIPKVLQYYSAATEHDRSWYKAWHAWAVMNFEA
VLHYKHQNQARDEKKKLRHASGANITNATTAATTAATATTTASTEGSNSESEAESTENSP
TPSPLQKKVTEDLSKTLLMYTVPAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVN
EALVEGVKAIQIDTWLQVIPQLIARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVAS
KSTTTARHNAANKILKNMCEHSNTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFG
ERNVKGMFEVLEPLHAMMERGPQTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQA
WDLYYHVFRRISKQLPQLTSLELQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSL
QVITSKQRPRKLTLMGSNGHEFVFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNL
SIQRYAVIPLSTNSGLIGWVPHCDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTL
MQKVEVFEHAVNNTAGDDLAKLLWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRH
PSNLMLDRLSGKILHIDFGDCFEVAMTREKFPEKIPFRLTRMLTNAMEVTGLDGNYRITC
HTVMEVLREHKDSVMAVLEAFVYDPLLNWRLMDTNTKGNKRSRTRTDSYSAGQSVEILDG
VELGEPAHKKTGTTVPESIHSFIGDGLVKPEALNKKAIQIINRVRDKLTGRDFSHDDTLD
VPTQVELLIKQATSHENLCQCYIGWCPFW

Sequence of entity 2 (C, D), FASTA

>5ZCS_2 Target of rapamycin complex subunit LST8 (chains C, D)
MNTSPGTVGSDPVILATAGYDHTVRFWQAHSGICTRTVQHQDSQVNALEVTPDRSMIAAA
GYQHIRMYDLNSNNPNPIISYDGVNKNIASVGFHEDGRWMYTGGEDCTARIWDLRSRNLQ
CQRIFQVNAPINCVCLHPNQAELIVGDQSGAIHIWDLKTDHNEQLIPEPEVSITSAHIDP
DASYMAAVNSTGNCYVWNLTGGIGDEVTQLIPKTKIPAHTRYALQCRFSPDSTLLATCSA
DQTCKIWRTSNFSLMTELSIKSGNPGESSRGWMWGCAFSGDSQYIVTASSDNLARLWCVE
TGEIKREYGGHQKAVVCLAFNDSVLG

Sequence of entity 3 (E, F), FASTA

>5ZCS_3 Rapamycin-insensitive companion of mTOR (chains E, F)
MAAIGRGRSLKNLRVRGRNDSGEENVPLDLTREPSDNLREILQNVARLQGVSNMRKLGHL
NNFTKLLCDIGHSEEKLGFHYEDIIICLRLALLNEAKEVRAAGLRALRYLIQDSSILQKV
LKLKVDYLIARCIDIQQSNEVERTQALRLVRKMITVNASLFPSSVTNSLIAVGNDGLQER
DRMVRACIAIICELALQNPEVVALRGGLNTILKNVIDCQLSRINEALITTILHLLNHPKT
RQYVRADVELERILAPYTDFHYRHSPDTAEGQLKEDREARFLASKMGIIATFRSWAGIIN
LCKPGNSGIQSLIGVLCIPNMEIRRGLLEVLYDIFRLPLPVVTEEFIEALLSVDPGRFQD
SWRLSDGFVAAEAKTILPHRARSRPDLMDNYLALILSAFIRNGLLEGLVEVITNSDDHIS
VRATILLGELLHMANTILPHSHSHHLHCLPTLMNMAASFDIPKEKRLRASAALNCLKRFH
EMKKRGPKPYSLHLDHIIQKAIATHQKRDQYLRVQKDIFILKDTEEALLINLRDSQVLQH
KENLEWNWNLIGTILKWPNVNLRNYKDEQLHRFVRRLLYFYKPSSKLYANLDLDFAKAKQ
LTVVGCQFTEFLLESEEDGQGYLEDLVKDIVQWLNASSGMKPERSLQNNGLLTTLSQHYF
LFIGTLSCHPHGVKMLEKCSVFQCLLNLCSLKNQDHLLKLTVSSLDYSRDGLARVILSKI
LTAATDACRLYATKHLRVLLRANVEFFNNWGIELLVTQLHDKNKTISSEALDILDEACED
KANLHALIQMKPALSHLGDKGLLLLLRFLSIPKGFSYLNERGYVAKQLEKWHREYNSKYV
DLIEEQLNEALTTYRKPVDGDNYVRRSNQRLQRPHVYLPIHLYGQLVHHKTGCHLLEVQN
IITELCRNVRTPDLDKWEEIKKLKASLWALGNIGSSNWGLNLLQEENVIPDILKLAKQCE
VLSIRGTCVYVLGLIAKTKQGCDILKCHNWDAVRHSRKHLWPVVPDDVEQLCNELSSIXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXX

Sequence of entity 4 (G, H), FASTA

>5ZCS_4 Target of rapamycin complex 2 subunit MAPKAP1 (chains G, H)
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXILSVRLEQCPLQLNNPFNEYSKFDGKGHVGTTATKKIDVY
LPLHSSQDRLLPMTVVTMASARVQDLIGLICWQYTSEGREPKLNDNVSAYCLHIAEDDGE
VDTDFPPLDSNEPIHKFGFSTLALVEKYSSPGLTSKESLFVRINAAHGFSLIQVDNTKVT
MKEILLKAVKRRKGSQKVSGPQYRLEKQSEPNVAVDLDSTLESQSAWEFCLVRENSSRAD
GVFEEDSQIDIATVQDMLSSHHYKSFKVSMIHRLRFTTDVQLGISGDKVEIDPVTNQKAS
TKFWIKQKPISIDSDLLCACDLAEEKSPSHAIFKLTYLSNHDYKHLYFESDAATVNEIVL
KVNYILESRASTARADYFAQKQRKLNRRTSFSFQKEKKSGQQ

Primary citation

Cryo-EM structure of human mTOR complex 2. Chen, X., Liu, M., Tian, Y. et al. Cell Res (2018) 28:518-528. DOI 10.1038/s41422-018-0029-3 · PubMed

Other PDB entries of the same protein (UniProt P42345 (AlphaFold model), which also has an AlphaFold model), best resolution first:

4DRI 1.45 Å, Co-crystal structure of the PPIase domain of FKBP51, Rapamycin and the FRB fragment of…
9DBO 1.55 Å, Crystal structure of a synthetic Fab (R3E9) in complex with the FRB domain of mTOR
5GPG 1.67 Å, Co-crystal structure of the FK506 binding domain of human FKBP25, Rapamycin and the FRB…
5WBH 1.75 Å, Structure of the FRB domain of mTOR bound to a substrate recruitment peptide of S6K1
4DRJ 1.8 Å, o-crystal structure of the PPIase domain of FKBP52, Rapamycin and the FRB fragment of mTOR
3FAP 1.85 Å, Atomic structures of the rapamycin analogs in complex with both human FKBP12 and frb…
8PPZ 1.85 Å, Co-crystal structure of FKBP12, compound 7 and the FRB fragment of mTOR
8XI9 1.85 Å, Crystal structure of FRB-FKBP fusion protein in complex with rapamycin
9PIW 1.9 Å, Crystal structure of a synthetic Fab (1A) in complex with the FRB domain of mTOR
9DL0 2.0 Å, Crystal structure of a synthetic Fab (R3H8) in complex with the FRB domain of mTOR
9PJ1 2.05 Å, Crystal structure of a synthetic Fab (4R) in complex with the ternary assembly of…
9PIU 2.18 Å, Crystal structure of a synthetic Fab (2C) in complex with the FRB domain of mTOR

Browse structure collections

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