9OTI: GATOR2 complex

GATOR2 complex bound to arginine sensor CASTOR1. Determined by electron microscopy at 3.5 Å resolution. Released 1 Oct 2025.

Method: Electron microscopy
Resolution: 3.5 Å
Organism: Homo sapiens
Chains: 18
Atoms: 43,325
Mol. weight: 1175.09 kDa
Ligands: ZN
Released: 1 Oct 2025

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

Secondary structure: helices and β-sheets

9OTI contains 250 α-helices and 390 β-strands across 18 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: 32 helices, 13 β-strands

Element	Residues	Length	Sheet
β-strand	356	1	1
β-strand	359	1	1
β-strand	365-370	6	1
β-strand	373-378	6	1
α-helix	390-399	10
α-helix	409-411	3
α-helix	412-415	4
α-helix	421-439	19
α-helix	452-454	3
α-helix	455-461	7
α-helix	488-497	10
α-helix	509-518	10
α-helix	521-530	10
α-helix	534-546	13
α-helix	553-561	9
α-helix	571-578	8
α-helix	580-582	3
α-helix	586-596	11
α-helix	598	1
α-helix	603-606	4
α-helix	613-623	11
α-helix	626-643	18
α-helix	646-648	3
α-helix	649-652	4
α-helix	656-669	14
α-helix	672-679	8
α-helix	685-689	5
α-helix	691-706	16
α-helix	710-723	14
α-helix	728-731	4
β-strand	734-736	3	2
β-strand	743	1	2
β-strand	774	1	3
α-helix	780	1
β-strand	781	1	3
α-helix	782-783	2
β-strand	784	1	4
α-helix	785	1
β-strand	791	1	4
α-helix	820-822	3
β-strand	824-827	4	5
β-strand	832	1	6
β-strand	835	1	5
α-helix	836-845	10
α-helix	858-862	5

Chain B: 32 helices, 40 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	7
β-strand	17-21	5	7
β-strand	25-30	6	7
β-strand	51-57	7	7
β-strand	63-65	3	8
β-strand	68	1	8
β-strand	78-82	5	8
β-strand	87-90	4	8
β-strand	107-108	2	8
β-strand	116-120	5	9
β-strand	130-133	4	9
β-strand	140-144	5	9
β-strand	179-183	5	9
β-strand	189	1	10
β-strand	192	1	10
β-strand	199-203	5	10
β-strand	207	1	11
β-strand	208-212	5	10
β-strand	220	1	10
β-strand	223	1	11
β-strand	228-232	5	12
β-strand	239-244	6	12
β-strand	247-252	6	12
α-helix	253-255	3
β-strand	256	1	13
β-strand	258	1	13
β-strand	261-262	2	12
α-helix	269	1
β-strand	270-274	5	14
β-strand	283-287	5	14
β-strand	293-296	4	14
β-strand	315-318	4	14
β-strand	327-330	4	15
β-strand	337-341	5	15
β-strand	348-351	4	15
α-helix	354-356	3
β-strand	367-370	4	16
β-strand	373-376	4	16
α-helix	390-399	10
α-helix	409-411	3
α-helix	412-415	4
α-helix	421-437	17
α-helix	452-454	3
α-helix	455-459	5
α-helix	488-496	9
α-helix	509-518	10
α-helix	521-530	10
α-helix	534-546	13
α-helix	553-561	9
α-helix	571-579	9
α-helix	580-582	3
α-helix	586-596	11
α-helix	598	1
α-helix	603-606	4
α-helix	613-623	11
α-helix	626-643	18
α-helix	646-648	3
α-helix	649-652	4
α-helix	656-668	13
α-helix	672-678	7
α-helix	685-688	4
α-helix	691-706	16
α-helix	710-723	14
α-helix	725-727	3
β-strand	784	1	17
β-strand	791	1	17
α-helix	820-822	3
β-strand	824	1	18
β-strand	825	1	19
β-strand	835	1	18
α-helix	836-845	10
α-helix	858-862	5

Chain C: 11 helices, 10 β-strands

Element	Residues	Length	Sheet
β-strand	344-345	2	20
β-strand	351-352	2	20
β-strand	400	1	20
α-helix	410-417	8
α-helix	426-440	15
α-helix	443-456	14
α-helix	634-646	13
α-helix	650-660	11
α-helix	662-665	4
α-helix	670-686	17
α-helix	690-699	10
α-helix	703-706	4
α-helix	707-709	3
β-strand	714	1	6
β-strand	716-719	4	2
β-strand	724-725	2	2
β-strand	742	1	21
β-strand	749	1	21
β-strand	753-756	4	5
β-strand	763-765	3	5
α-helix	766-773	8

Chain D: 16 helices, 7 β-strands

Element	Residues	Length	Sheet
α-helix	534-538	5
β-strand	541-545	5	22
β-strand	549-553	5	22
β-strand	646-651	6	22
α-helix	653-655	3
α-helix	660-665	6
α-helix	673-687	15
α-helix	690-702	13
α-helix	705-707	3
α-helix	708-710	3
α-helix	723-727	5
α-helix	728-738	11
α-helix	742-755	14
α-helix	838-847	10
α-helix	853-855	3
α-helix	856-873	18
α-helix	876-883	8
β-strand	923	1	23
β-strand	930	1	23
β-strand	935-937	3	19
β-strand	944-946	3	19
α-helix	947-954	8
α-helix	968-972	5

Chain E: 3 helices, 27 β-strands

Element	Residues	Length	Sheet
β-strand	3-6	4	1
β-strand	15-20	6	24
β-strand	27-31	5	24
β-strand	35-40	6	24
β-strand	48-54	7	24
β-strand	60-65	6	25
α-helix	68-70	3
β-strand	73-78	6	25
β-strand	83-88	6	25
β-strand	103-108	6	25
β-strand	116-121	6	26
α-helix	122-123	2
β-strand	125	1	27
β-strand	127	1	27
β-strand	129-134	6	26
β-strand	138-143	6	26
β-strand	154-160	7	26
β-strand	168-170	3	28
β-strand	181-184	4	28
β-strand	197-203	7	28
β-strand	208-215	8	28
α-helix	221-222	2
β-strand	224-227	4	29
β-strand	236-241	6	29
β-strand	245-252	8	29
β-strand	266-274	9	29
β-strand	281-286	6	30
β-strand	292-297	6	30
β-strand	302-307	6	30
β-strand	313-318	6	30

Chain F: 5 helices, 28 β-strands

Element	Residues	Length	Sheet
β-strand	5-7	3	16
β-strand	15-20	6	31
β-strand	27-31	5	31
β-strand	35-39	5	31
β-strand	49	1	31
β-strand	62-65	4	32
α-helix	68-70	3
β-strand	74-77	4	32
β-strand	83-86	4	32
β-strand	107-109	3	32
α-helix	115-116	2
β-strand	120-121	2	33
α-helix	122-123	2
α-helix	124-126	3
β-strand	129-131	3	33
β-strand	133	1	33
β-strand	139-143	5	33
β-strand	154-159	6	33
β-strand	167-170	4	34
β-strand	181-185	5	34
β-strand	197-201	5	34
β-strand	202-203	2	35
β-strand	208-209	2	35
β-strand	225-227	3	36
α-helix	228-229	2
β-strand	236-240	5	36
β-strand	246-250	5	36
β-strand	281	1	37
β-strand	285-286	2	38
β-strand	293-295	3	38
β-strand	297	1	37
β-strand	301-306	6	38
β-strand	314-320	7	38

Chain G: 5 helices, 26 β-strands

Element	Residues	Length	Sheet
β-strand	17-20	4	39
β-strand	26-30	5	39
β-strand	36-40	5	39
β-strand	48-53	6	39
β-strand	60-65	6	40
α-helix	68-70	3
β-strand	73-78	6	40
β-strand	83-89	7	40
β-strand	102-109	8	40
β-strand	116-121	6	41
α-helix	122-123	2
α-helix	124-126	3
β-strand	129-134	6	41
β-strand	138-143	6	41
β-strand	154-160	7	41
β-strand	165-170	6	42
β-strand	181-186	6	42
β-strand	197-203	7	42
β-strand	208-214	7	42
β-strand	224-227	4	43
α-helix	228-229	2
β-strand	236-241	6	43
β-strand	245-250	6	43
β-strand	251-252	2	44
α-helix	263-265	3
β-strand	266-267	2	44
β-strand	270-274	5	43
β-strand	281-286	6	45
β-strand	293-297	5	45
β-strand	302-307	6	45
β-strand	313-319	7	45

Chain H: 4 helices, 26 β-strands

Element	Residues	Length	Sheet
β-strand	5-9	5	22
β-strand	16-21	6	46
β-strand	27-32	6	46
β-strand	37-41	5	46
β-strand	48-53	6	46
β-strand	60-65	6	47
α-helix	66-67	2
α-helix	68-70	3
β-strand	73-78	6	47
β-strand	83-89	7	47
β-strand	92-99	8	47
β-strand	106-111	6	48
α-helix	112-113	2
β-strand	119-124	6	48
β-strand	129-136	8	48
β-strand	139-142	4	48
β-strand	145	1	48
β-strand	153-158	6	49
β-strand	181-186	6	49
β-strand	191-196	6	49
β-strand	202-208	7	49
β-strand	215-220	6	50
α-helix	221-223	3
β-strand	229-235	7	50
β-strand	239-245	7	50
β-strand	256-260	5	50
β-strand	265-270	6	51
β-strand	276-281	6	51
β-strand	286-290	5	51
β-strand	298-299	2	51

10 more chain groups are not listed. Open the entry in the viewer and use the sequence panel to see them.

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
GATOR2 complex protein MIOS	A, B, L, T	protein	875	Homo sapiens	Q9NXC5 (AlphaFold model)
GATOR complex protein WDR24	C, M	protein	790	Homo sapiens	Q96S15 (AlphaFold model)
GATOR complex protein WDR59	D, N	protein	974	Homo sapiens	Q6PJI9 (AlphaFold model)
Nucleoporin SEH1	E, F, G, O, P, Q	protein	360	Homo sapiens	Q96EE3 (AlphaFold model)
Protein SEC13 homolog	H, R	protein	322	Homo sapiens	P55735
Cytosolic arginine sensor for mTORC1 subunit 1	U, V	protein	329	Homo sapiens	Q8WTX7

Sequence of entity 1 (A, B, L, T), FASTA

>9OTI_1 GATOR2 complex protein MIOS (chains A, B, L, T)
MSGTKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDT
PYMKCVAWYLNYDPECLLAVGQANGRVVLTSLGQDHNSKFKDLIGKEFVPKHARQCNTLA
WNPLDSNWLAAGLDKHRADFSVLIWDICSKYTPDIVPMEKVKLSAGETETTLLVTKPLYE
LGQNDACLSLCWLPRDQKLLLAGMHRNLAIFDLRNTSQKMFVNTKAVQGVTVDPYFHDRV
ASFYEGQVAIWDLRKFEKPVLTLTEQPKPLTKVAWCPTRTGLLATLTRDSNIIRLYDMQH
TPTPIGDETEPTIIERSVQPCDNYIASFAWHPTSQNRMIVVTPNRTMSDFTVFERISLAW
SPITSLMWACGRHLYECTEEENDNSLEKDIATKMRLRALSRYGLDTEQVWRNHILAGNED
PQLKSLWYTLHFMKQYTEDMDQKSPGNKGSLVYAGIKSIVKSSLGMVESSRHNWSGLDKQ
SDIQNLNEERILALQLCGWIKKGTDVDVGPFLNSLVQEGEWERAAAVALFNLDIRRAIQI
LNEGASSEKGDLNLNVVAMALSGYTDEKNSLWREMCSTLRLQLNNPYLCVMFAFLTSETG
SYDGVLYENKVAVRDRVAFACKFLSDTQLNRYIEKLTNEMKEAGNLEGILLTGLTKDGVD
LMESYVDRTGDVQTASYCMLQGSPLDVLKDERVQYWIENYRNLLDAWRFWHKRAEFDIHR
SKLDPSSKPLAQVFVSCNFCGKSISYSCSAVPHQGRGFSQYGVSGSPTKSKVTSCPGCRK
PLPRCALCLINMGTPVSSCPGGTKSDEKVDLSKDKKLAQFNNWFTWCHNCRHGGHAGHML
SWFRDHAECPVSACTCKCMQLDTTGNLVPAETVQP

Sequence of entity 2 (C, M), FASTA

>9OTI_2 GATOR complex protein WDR24 (chains C, M)
MEKMSRVTTALGGSVLTGRTMHCHLDAPANAISVCRDAAQVVVAGRSIFKIYAIEEEQFV
EKLNLRVGRKPSLNLSCADVVWHQMDENLLATAATNGVVVTWNLGRPSRNKQDQLFTEHK
RTVNKVCFHPTEAHVLLSGSQDGFMKCFDLRRKDSVSTFSGQSESVRDVQFSIRDYFTFA
STFENGNVQLWDIRRPDRCERMFTAHNGPVFCCDWHPEDRGWLATGGRDKMVKVWDMTTH
RAKEMHCVQTIASVARVKWRPECRHHLATCSMMVDHNIYVWDVRRPFVPAAMFEEHRDVT
TGIAWRHPHDPSFLLSGSKDSSLCQHLFRDASQPVERANPEGLCYGLFGDLAFAAKESLV
AAESGRKPYTGDRRHPIFFKRKLDPAEPFAGLASSALSVFETEPGGGGMRWFVDTAERYA
LAGRPLAELCDHNAKVARELGRNQVAQTWTMLRIIYCSPGLVPTANLNHSVGKGGSCGLP
LMNSFNLKDMAPGLGSETRLDRSKGDARSDTVLLDSSATLITNEDNEETEGSDVPADYLL
GDVEGEEDELYLLDPEHAHPEDPECVLPQEAFPLRHEIVDTPPGPEHLQDKADSPHVSGS
EADVASLAPVDSSFSLLSVSHALYDSRLPPDFFGVLVRDMLHFYAEQGDVQMAVSVLIVL
GERVRKDIDEQTQEHWYTSYIDLLQRFRLWNVSNEVVKLSTSRAVSCLNQASTTLHVNCS
HCKRPMSSRGWVCDRCHRCASMCAVCHHVVKGLFVWCQGCSHGGHLQHIMKWLEGSSHCP
AGCGHLCEYS

Sequence of entity 3 (D, N), FASTA

>9OTI_3 GATOR complex protein WDR59 (chains D, N)
MAARWSSENVVVEFRDSQATAMSVDCLGQHAVLSGRRFLYIVNLDAPFEGHRKISRQSKW
DIGAVQWNPHDSFAHYFAASSNQRVDLYKWKDGSGEVGTTLQGHTRVISDLDWAVFEPDL
LVTSSVDTYIYIWDIKDTRKPTVALSAVAGASQVKWNKKNANCLATSHDGDVRIWDKRKP
STAVEYLAAHLSKIHGLDWHPDSEHILATSSQDNSVKFWDYRQPRKYLNILPCQVPVWKA
RYTPFSNGLVTVMVPQLRRENSLLLWNVFDLNTPVHTFVGHDDVVLEFQWRKQKEGSKDY
QLVTWSRDQTLRMWRVDSQMQRLCANDILDGVDEFIESISLLPEPEKTLHTEDTDHQHTA
SHGEEEALKEDPPRNLLEERKSDQLGLPQTLQQEFSLINVQIRNVNVEMDAADRSCTVSV
HCSNHRVKMLVKFPAQYPNNAAPSFQFINPTTITSTMKAKLLKILKDTALQKVKRGQSCL
EPCLRQLVSCLESFVNQEDSASSNPFALPNSVTPPLPTFARVTTAYGSYQDANIPFPRTS
GARFCGAGYLVYFTRPMTMHRAVSPTEPTPRSLSALSAYHTGLIAPMKIRTEAPGNLRLY
SGSPTRSEKEQVSISSFYYKERKSRRWKSKREGSDSGNRQIKAAGKVIIQDIACLLPVHK
SLGELYILNVNDIQETCQKNAASALLVGRKDLVQVWSLATVATDLCLGPKSDPDLETPWA
RHPFGRQLLESLLAHYCRLRDVQTLAMLCSVFEAQSRPQGLPNPFGPFPNRSSNLVVSHS
RYPSFTSSGSCSSMSDPGLNTGGWNIAGREAEHLSSPWGESSPEELRFGSLTYSDPRERE
RDQHDKNKRLLDPANTQQFDDFKKCYGEILYRWGLREKRAEVLKFVSCPPDPHKGIEFGV
YCSHCRSEVRGTQCAICKGFTFQCAICHVAVRGSSNFCLTCGHGGHTSHMMEWFRTQEVC
PTGCGCHCLLESTF

Sequence of entity 4 (E, F, G, O, P, Q), FASTA

>9OTI_4 Nucleoporin SEH1 (chains E, F, G, O, P, Q)
MFVARSIAADHKDLIHDVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHSGSV
WRVTWAHPEFGQVLASCSFDRTAAVWEEIVGESNDKLRGQSHWVKRTTLVDSRTSVTDVK
FAPKHMGLMLATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSP
MIAVGSDDSSPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAI
ATKDVRIFTLKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDG
CVRLWKANYMDNWKCTGILKGNGSPVNGSSQQGTSNPSLGSTIPSLQNSLNGSSAGRKHS

Sequence of entity 5 (H, R), FASTA

>9OTI_5 Protein SEC13 homolog (chains H, R)
MVSVINTVDTSHEDMIHDAQMDYYGTRLATCSSDRSVKIFDVRNGGQILIADLRGHEGPV
WQVAWAHPMYGNILASCSYDRKVIIWREENGTWEKSHEHAGHDSSVNSVCWAPHDYGLIL
ACGSSDGAISLLTYTGEGQWEVKKINNAHTIGCNAVSWAPAVVPGSLIDHPSGQKPNYIK
RFASGGCDNLIKLWKEEEDGQWKEEQKLEAHSDWVRDVAWAPSIGLPTSTIASCSQDGRV
FIWTCDDASSNTWSPKLLHKFNDVVWHVSWSITANILAVSGGDNKVTLWKESVDGQWVCI
SDVNKGQGSVSASVTEGQQNEQ

Sequence of entity 6 (U, V), FASTA

>9OTI_6 Cytosolic arginine sensor for mTORC1 subunit 1 (chains U, V)
MELHILEHRVRVLSVARPGLWLYTHPLIKLLFLPRRSRCKFFSLTETPEDYTLMVDEEGF
KELPPSEFLQVAEATWLVLNVSSHSGAAVQAAGVTKIARSVIAPLAEHHVAVLMLSTYQT
DFILVREQDLSVVIHTLAQEFDIYREVGGEPVPVTRDDSSNGFPRTQHGPSPTVHPIQSP
QNRFCVLTLDPETLPAIATTLIDVLFYSHSTPKEAASSSPEPSSITFFAFSLIEGYISIV
MDAETQKKFPSDLLLTSSSGELWRMVRIGGQPLGFDECGIVAQIAGPLAAADISAYYIST
FNFAHALVPEDGIGSVIEVLQRRQEGLAS

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	29

Primary citation

Structural basis for mTORC1 regulation by the CASTOR1-GATOR2 complex. Jansen, R.M., Maghe, C., Tapia, K. et al. Nat Struct Mol Biol (2025) 32:1980-1988. DOI 10.1038/s41594-025-01635-0 · PubMed

Other PDB entries of the same protein (UniProt Q9NXC5 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9DX1 3.36 Å, Human GATOR2 complex - Sestrin2 bound state
9DX2 3.4 Å, Human GATOR2 complex - CASTOR1 bound state
9LWF 3.41 Å, Cryo-EM structure of dual sensor bound GATOR2 complex
9DX0 3.47 Å, Human GATOR2 complex - apo state
9LVK 3.59 Å, Cryo-EM structure of CASTOR1 bound human GATOR2 complex
7UHY 3.66 Å, Human GATOR2 complex
9LVJ 3.82 Å, Cryo-EM structure of Sestrin2 bound human GATOR2 complex

Browse structure collections

About this viewer

MolViewer shows 9OTI directly in your browser with nothing to install. Switch between cartoon, ball-and-stick, spacefill and surface views, color by chain, secondary structure or B-factor, measure distances, angles and dihedrals, and share or embed the view.