9ZBJ: Human apo mTORC2

Cryo-EM structure of human apo mTORC2. Determined by electron microscopy at 3.2 Å resolution. Released 15 Apr 2026.

Method: Electron microscopy
Resolution: 3.2 Å
Organism: Homo sapiens
Chains: 4
Atoms: 28,107
Mol. weight: 507.58 kDa
Ligands: ZN
Released: 15 Apr 2026

Explore 9ZBJ in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9ZBJ contains 227 α-helices and 47 β-strands across 4 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.

Chain A: 139 helices, 12 β-strands

Element	Residues	Length	Sheet
α-helix	204-223	20
α-helix	234-244	11
α-helix	258-273	16
α-helix	279-289	11
α-helix	361-367	7
α-helix	372-381	10
α-helix	386-394	9
α-helix	396-400	5
α-helix	404-408	5
α-helix	414-422	9
α-helix	427-429	3
α-helix	430	1
α-helix	431-435	5
α-helix	436-443	8
α-helix	445-448	4
α-helix	452-459	8
α-helix	463-466	4
α-helix	478-493	16
α-helix	495-497	3
α-helix	498-511	14
α-helix	515-527	13
α-helix	529-531	3
α-helix	532-546	15
α-helix	579-591	13
α-helix	600-609	10
α-helix	616-628	13
α-helix	631-635	5
α-helix	647-664	18
α-helix	669-678	10
α-helix	681-683	3
α-helix	684-687	4
α-helix	690-699	10
α-helix	705-718	14
α-helix	723-743	21
α-helix	747-763	17
α-helix	765-768	4
α-helix	769-771	3
α-helix	772-783	12
α-helix	792-809	18
α-helix	811-815	5
α-helix	817-828	12
α-helix	834-851	18
α-helix	856-860	5
α-helix	864-874	11
α-helix	878-891	14
α-helix	896-903	8
α-helix	929-936	8
α-helix	943-959	17
α-helix	962-964	3
α-helix	965-982	18
α-helix	983-989	7
α-helix	994-1003	10
α-helix	1006-1022	17
α-helix	1028-1030	3
α-helix	1031-1041	11
α-helix	1048-1063	16
α-helix	1064-1069	6
α-helix	1071-1084	14
α-helix	1088-1090	3
α-helix	1091-1103	13
α-helix	1104-1107	4
α-helix	1111-1113	3
α-helix	1115-1123	9
α-helix	1129-1145	17
α-helix	1152-1165	14
α-helix	1170-1184	15
α-helix	1185-1191	7
α-helix	1192-1202	11
α-helix	1207-1218	12
α-helix	1229-1240	12
α-helix	1265-1269	5
α-helix	1277-1294	18
α-helix	1298-1301	4
α-helix	1304-1309	6
α-helix	1311-1327	17
α-helix	1331-1347	17
α-helix	1351-1366	16
α-helix	1380-1389	10
α-helix	1393-1406	14
α-helix	1410-1422	13
α-helix	1426-1439	14
α-helix	1447-1453	7
α-helix	1456-1466	11
α-helix	1474-1487	14
α-helix	1490-1500	11
α-helix	1506-1523	18
α-helix	1527-1533	7
α-helix	1541-1553	13
α-helix	1557-1579	23
α-helix	1583-1607	25
α-helix	1612-1624	13
α-helix	1630-1640	11
α-helix	1646-1648	3
α-helix	1650-1663	14
α-helix	1666-1677	12
α-helix	1681-1683	3
α-helix	1688-1690	3
α-helix	1694-1706	13
α-helix	1711-1730	20
α-helix	1737-1762	26
α-helix	1768-1782	15
α-helix	1787-1813	27
α-helix	1872-1875	4
α-helix	1878-1880	3
α-helix	1882-1894	13
α-helix	1900-1913	14
α-helix	1917-1929	13
α-helix	1933-1937	5
α-helix	1939-1945	7
α-helix	1951-1967	17
α-helix	1970-1980	11
α-helix	1985-2020	36
α-helix	2023-2041	19
α-helix	2045-2060	16
α-helix	2065-2091	27
α-helix	2094-2114	21
α-helix	2115-2117	3
β-strand	2120-2122	3	1
α-helix	2123-2126	4
α-helix	2128-2132	5
α-helix	2138	1
β-strand	2139	1	2
α-helix	2140	1
α-helix	2149-2151	3
β-strand	2152	1	2
β-strand	2153-2156	4	1
β-strand	2159-2162	4	1
β-strand	2170-2176	7	1
β-strand	2181-2188	8	1
α-helix	2193-2211	19
α-helix	2213-2218	6
β-strand	2227-2229	3	1
β-strand	2234-2238	5	1
β-strand	2243-2245	3	3
α-helix	2246-2257	12
α-helix	2263-2271	9
α-helix	2280-2292	13
α-helix	2298-2306	9
α-helix	2310-2334	25
α-helix	2341-2343	3
β-strand	2344-2347	4	3
β-strand	2353-2355	3	3
α-helix	2365-2367	3
α-helix	2381-2386	6
α-helix	2395-2409	15
α-helix	2411-2422	12
α-helix	2429-2432	4
α-helix	2493-2509	17
α-helix	2521-2533	13
α-helix	2535-2538	4
α-helix	2543-2545	3

Chain B: 1 helix, 28 β-strands

Element	Residues	Length	Sheet
β-strand	14-19	6	4
β-strand	23-27	5	4
β-strand	34-39	6	4
β-strand	47-50	4	5
β-strand	56-60	5	5
β-strand	65-69	5	5
β-strand	79-81	3	5
β-strand	88-93	6	6
β-strand	99-104	6	6
β-strand	108-113	6	6
β-strand	124-126	3	6
β-strand	131-136	6	7
β-strand	142-147	6	7
β-strand	152-156	5	7
β-strand	162-165	4	7
β-strand	173-178	6	8
β-strand	184-189	6	8
β-strand	193-201	9	8
α-helix	207	1
β-strand	208-217	10	8
β-strand	223-228	6	9
β-strand	234-239	6	9
β-strand	243-248	6	9
β-strand	254-259	6	9
β-strand	273-277	5	10
β-strand	284-289	6	10
β-strand	293-298	6	10
β-strand	304-308	5	10
β-strand	315-321	7	4

Chain C: 80 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	34-46	13
α-helix	53-70	18
α-helix	74-77	4
α-helix	81-91	11
α-helix	97-110	14
α-helix	114-122	9
α-helix	126-134	9
α-helix	140-156	17
α-helix	164-174	11
α-helix	184-197	14
α-helix	199-205	7
α-helix	208-216	9
α-helix	221-236	16
α-helix	238-241	4
α-helix	250-254	5
α-helix	255-258	4
α-helix	273-292	20
α-helix	295-301	7
α-helix	308-317	10
α-helix	321-334	14
α-helix	338-342	5
α-helix	346-352	7
α-helix	360-362	3
α-helix	368-376	9
α-helix	378-379	2
β-strand	386	1	11
α-helix	387-402	16
α-helix	404-413	10
α-helix	417-436	20
α-helix	440-443	4
α-helix	446-448	3
α-helix	450-456	7
α-helix	463-485	23
α-helix	492-508	17
α-helix	515-517	3
α-helix	525-534	10
α-helix	548-556	9
α-helix	568-581	14
α-helix	583-585	3
β-strand	592	1	12
α-helix	598-613	16
α-helix	620-638	19
α-helix	642-647	6
α-helix	648-651	4
β-strand	654	1	12
α-helix	655-658	4
α-helix	659-667	9
α-helix	670-678	9
α-helix	681-688	8
α-helix	695-704	10
α-helix	712-720	9
α-helix	726-741	16
α-helix	747-757	11
α-helix	758-760	3
α-helix	764-777	14
α-helix	781-790	10
α-helix	795-797	3
α-helix	799-806	8
α-helix	807-810	4
α-helix	812-820	9
α-helix	823-830	8
α-helix	831-835	5
α-helix	836-851	16
β-strand	877	1	11
α-helix	878-880	3
α-helix	882-886	5
α-helix	890-898	9
α-helix	901-910	10
α-helix	917-933	17
α-helix	937-946	10
α-helix	949-958	10
α-helix	962-975	14
α-helix	979-986	8
β-strand	990-992	3	13
α-helix	1002-1004	3
β-strand	1428-1430	3	13
α-helix	1433-1435	3
α-helix	1480-1486	7
α-helix	1524-1526	3
α-helix	1531-1535	5
α-helix	1608-1624	17
α-helix	1629-1642	14
α-helix	1644-1647	4
α-helix	1650-1662	13
α-helix	1667-1675	9
α-helix	1683-1693	11

Chain D: 7 helices, 1 β-strand

Element	Residues	Length	Sheet
α-helix	7-20	14
α-helix	26-30	5
α-helix	71-73	3
α-helix	85-98	14
β-strand	106	1	5
α-helix	109-111	3
α-helix	117-122	6
α-helix	140-147	8

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Serine/threonine-protein kinase mTOR	A	protein	2347	Homo sapiens	P42345 (AlphaFold model)
Target of rapamycin complex subunit LST8	B	protein	317	Homo sapiens	Q9BVC4 (AlphaFold model)
Rapamycin-insensitive companion of mTOR	C	protein	1672	Homo sapiens	Q6R327 (AlphaFold model)
Target of rapamycin complex 2 subunit MAPKAP1	D	protein	148	Homo sapiens	Q9BPZ7 (AlphaFold model)

Sequence of entity 1 (A), FASTA

>9ZBJ_1 Serine/threonine-protein kinase mTOR (chains A)
QAIREGAVAALRACLILTTQREPKEMQKPQWYRHTFEEAEKGFDETLAKEKGMNRDDRIH
GALLILNELVRISSMEGERLREEMEEITQQQLVHDKYCKDLMGFGTKPRHITPFTSFQAV
QPQQSNALVGLLGYSSHQGLMGFGTSPSPAKSTLVESRCCRDLMEEKFDQVCQWVLKCRN
SKNSLIQMTILNLLPRLAAFRPSAFTDTQYLQDTMNHVLSCVKKEKERTAAFQALGLLSV
AVRSEFKVYLPRVLDIIRAALPPKDFAHKRQKAMQVDATVFTCISMLARAMGPGIQQDIK
ELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLLKMLSLVLMHKPLRHPGMPKGLA
HQLASPGLTTLPEASDVGSITLALRTLGSFEFEGHSLTQFVRHCADHFLNSEHKEIRMEA
ARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLLVVGITDPDPDIRYCVLASLDER
FDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSSMNPAFVMPFLRKMLIQILTELE
HSGIGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALILKLKDPDPDPNPGVINNVLATI
GELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVALWTLGQLVASTGYVVEPYRKYP
TLLEVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHKVNIGMIDQSRDASAVSLSESKS
SQDSSDYSTSEMLVNMGNLPLDEFYPAVSMVALMRIFRDQSLSHHHTMVVQAITFIFKSL
GLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLVSFVKSHIRPYMDEIVTLMREFW
VMNTSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLRVFMHDNSPGRIVSIKLLAAIQL
FGANLDDYLHLLLPPIVKLFDAPEAPLPSRKAALETVDRLTESLDFTDYASRIIHPIVRT
LDQSPELRSTAMDTLSSLVFQLGKKYQIFIPMVNKVLVRHRINHQRYDVLICRIVKGYTL
ADEEEDPLIYQHRMLRSGQGDALASGPVETGPMKKLHVSTINLQKAWGAARRVSKDDWLE
WLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNAAFVSCWSELNEDQQDELIRSIE
LALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGIVLLGERAAKCRAYAKALHYKEL
EFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHFGELEIQATWYEKLHEWEDALVA
YDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEKWTLVNDETQAKMARMAAAAAWG
LGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLAQQCIDKARDLLDAELTAMAGES
YSRAYGAMVSCHMLSELEEVIQYKLVPERREIIRQIWWERLQGCQRIVEDWQKILMVRSL
VVSPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVDPSRQLDHPLPTVHPQVTYAYMK
NMWKSARKIDAFQHMQHFVQTMQQQAQHAIATEDQQHKQELHKLMARCFLKLGEWQLNLQ
GINESTIPKVLQYYSAATEHDRSWYKAWHAWAVMNFEAVLHYKHQNQARDEKKKLRHASG
ANITNATTAATTAATATTTASTEGSNSESEAESTENSPTPSPLQKKVTEDLSKTLLMYTV
PAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVNEALVEGVKAIQIDTWLQVIPQL
IARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVASKSTTTARHNAANKILKNMCEHS
NTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFGERNVKGMFEVLEPLHAMMERGP
QTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQAWDLYYHVFRRISKQLPQLTSLE
LQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSLQVITSKQRPRKLTLMGSNGHEF
VFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNLSIQRYAVIPLSTNSGLIGWVPH
CDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTLMQKVEVFEHAVNNTAGDDLAKL
LWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRHPSNLMLDRLSGKILHIDFGDCF
EVAMTREKFPEKIPFRLTRMLTNAMEVTGLDGNYRITCHTVMEVLREHKDSVMAVLEAFV
YDPLLNWRLMDTNTKGNKRSRTRTDSYSAGQSVEILDGVELGEPAHKKTGTTVPESIHSF
IGDGLVKPEALNKKAIQIINRVRDKLTGRDFSHDDTLDVPTQVELLIKQATSHENLCQCY
IGWCPFW

Sequence of entity 2 (B), FASTA

>9ZBJ_2 Target of rapamycin complex subunit LST8 (chains B)
TVGSDPVILATAGYDHTVRFWQAHSGICTRTVQHQDSQVNALEVTPDRSMIAAAGYQHIR
MYDLNSNNPNPIISYDGVNKNIASVGFHEDGRWMYTGGEDCTARIWDLRSRNLQCQRIFQ
VNAPINCVCLHPNQAELIVGDQSGAIHIWDLKTDHNEQLIPEPEVSITSAHIDPDASYMA
AVNSTGNCYVWNLTGGIGDEVTQLIPKTKIPAHTRYALQCRFSPDSTLLATCSADQTCKI
WRTSNFSLMTELSIKSGNPGESSRGWMWGCAFSGDSQYIVTASSDNLARLWCVETGEIKR
EYGGHQKAVVCLAFNDS

Sequence of entity 3 (C), FASTA

>9ZBJ_3 Rapamycin-insensitive companion of mTOR (chains C)
EENVPLDLTREPSDNLREILQNVARLQGVSNMRKLGHLNNFTKLLCDIGHSEEKLGFHYE
DIIICLRLALLNEAKEVRAAGLRALRYLIQDSSILQKVLKLKVDYLIARCIDIQQSNEVE
RTQALRLVRKMITVNASLFPSSVTNSLIAVGNDGLQERDRMVRACIAIICELALQNPEVV
ALRGGLNTILKNVIDCQLSRINEALITTILHLLNHPKTRQYVRADVELERILAPYTDFHY
RHSPDTAEGQLKEDREARFLASKMGIIATFRSWAGIINLCKPGNSGIQSLIGVLCIPNME
IRRGLLEVLYDIFRLPLPVVTEEFIEALLSVDPGRFQDSWRLSDGFVAAEAKTILPHRAR
SRPDLMDNYLALILSAFIRNGLLEGLVEVITNSDDHISVRATILLGELLHMANTILPHSH
SHHLHCLPTLMNMAASFDIPKEKRLRASAALNCLKRFHEMKKRGPKPYSLHLDHIIQKAI
ATHQKRDQYLRVQKDIFILKDTEEALLINLRDSQVLQHKENLEWNWNLIGTILKWPNVNL
RNYKDEQLHRFVRRLLYFYKPSSKLYANLDLDFAKAKQLTVVGCQFTEFLLESEEDGQGY
LEDLVKDIVQWLNASSGMKPERSLQNNGLLTTLSQHYFLFIGTLSCHPHGVKMLEKCSVF
QCLLNLCSLKNQDHLLKLTVSSLDYSRDGLARVILSKILTAATDACRLYATKHLRVLLRA
NVEFFNNWGIELLVTQLHDKNKTISSEALDILDEACEDKANLHALIQMKPALSHLGDKGL
LLLLRFLSIPKGFSYLNERGYVAKQLEKWHREYNSKYVDLIEEQLNEALTTYRKPVDGDN
YVRRSNQRLQRPHVYLPIHLYGQLVHHKTGCHLLEVQNIITELCRNVRTPDLDKWEEIKK
LKASLWALGNIGSSNWGLNLLQEENVIPDILKLAKQCEVLSIRGTCVYVLGLIAKTKQGC
DILKCHNWDAVRHSRKHLWPVVPDDVEQLCNELSSIPSTLSLNSESTSSRHNSESESVPS
SMFILEDDRFGSSSTSTFFLDINEDTEPTFYDRSGPIKDKNSFPFFASSKLVKNRILNSL
TLPNKKHRSSSDPKGGKLSSESKTSNRRIRTLTEPSVDFNHSDDFTPISTVQKTLQLETS
FMGNKHIEDTGSTPSIGENDLKFTKNFGTENHRENTSRERLVVESSTSSHMKIRSQSFNT
DTTTSGISSMSSSPSRETVGVDATTMDTDCGSMSTVVSTKTIKTSHYLTPQSNHLSLSKS
NSVSLVPPGSSHTLPRRAQSLKAPSIATIKSLADCNFSYTSSRDAFGYATLKRLQQQRMH
PSLSHSEALASPAKDVLFTDTITMKANSFESRLTPSRFMKALSYASLDKEDLLSPINQNT
LQRSSSVRSMVSSATYGGSDDYIGLALPVDINDIFQVKDIPYFQTKNIPPHDDRGARAFA
HDAGGLPSGTGGLVKNSFHLLRQQMSLTEIMNSIHSDASLFLESTEDTGLQEHTDDNCLY
CVCIEILGFQPSNQLSAICSHSDFQDIPYSDWCEQTIHNPLEVVPSKFSGISGCSDGVSQ
EGSASSTKSTELLLGVKTIPDDTPMCRILLRKEVLRLVINLSSSVSTKCHETGLLTIKEK
YPQTFDDICLYSEVSHLLSHCTFRLPCRRFIQELFQDVQFLQMHEEAEAVLA

Sequence of entity 4 (D), FASTA

>9ZBJ_4 Target of rapamycin complex 2 subunit MAPKAP1 (chains D)
MAFLDNPTIILAHIRQSHVTSDDTGMCEMVLIDHDVDLEKIHPPSMPGDSGSEIQGSNGE
TQGYVYAQSVDITSSWDFGIRRRSNTAQRLERLRKERQNQIKCKNIQWKERNSKQSAQEL
KSLFEKKSLKEKPPISGKQSILSVRLEQ

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	1

Primary citation

Structural basis for the recruitment and selective phosphorylation of Akt by mTORC2. Taylor, M.S., Chen, M., Hancock, M. et al. Science (2026) 391:eadv7111-eadv7111. DOI 10.1126/science.adv7111 · 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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