9DX0: Human GATOR2 complex - apo state

Human GATOR2 complex - apo state. Determined by electron microscopy at 3.47 Å resolution. Released 20 May 2026.

Method: Electron microscopy
Resolution: 3.47 Å
Organism: Homo sapiens
Chains: 4
Atoms: 27,053
Mol. weight: 424.54 kDa
Released: 20 May 2026

Explore 9DX0 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9DX0 contains 117 α-helices and 237 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: 37 helices, 66 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	1
β-strand	18-21	4	1
β-strand	25-32	8	1
α-helix	37-39	3
β-strand	42-46	5	1
β-strand	49-57	9	1
β-strand	63-68	6	2
β-strand	78-82	5	2
β-strand	87-90	4	2
β-strand	108	1	2
β-strand	116-121	6	3
β-strand	128-133	6	3
β-strand	140-146	7	3
β-strand	181-183	3	3
β-strand	187-192	6	4
β-strand	199-204	6	4
β-strand	207	1	5
β-strand	208-212	5	4
β-strand	223	1	5
β-strand	228-232	5	6
β-strand	239-244	6	6
β-strand	247-252	6	6
β-strand	261-262	2	6
α-helix	263-265	3
β-strand	270-275	6	7
β-strand	282-287	6	7
β-strand	293-298	6	7
β-strand	313-318	6	7
α-helix	324-325	2
β-strand	327-330	4	8
β-strand	337-341	5	8
β-strand	347-351	5	8
β-strand	356-359	4	9
β-strand	367-370	4	9
β-strand	373-376	4	9
α-helix	387-389	3
β-strand	390-391	2	9
β-strand	400-405	6	10
β-strand	411-416	6	10
β-strand	420-426	7	10
β-strand	432-439	8	10
β-strand	445-449	5	11
α-helix	453-455	3
β-strand	458-463	6	11
β-strand	468-475	8	11
β-strand	486-494	9	11
β-strand	501-506	6	12
α-helix	507-508	2
α-helix	509-511	3
β-strand	514-519	6	12
β-strand	523-528	6	12
β-strand	539-545	7	12
β-strand	550-555	6	13
β-strand	566-571	6	13
β-strand	582-588	7	13
β-strand	593-600	8	13
β-strand	609-612	4	14
β-strand	621-626	6	14
β-strand	630-637	8	14
α-helix	648-649	2
β-strand	651-659	9	14
β-strand	666-671	6	15
β-strand	678-682	5	15
β-strand	686-692	7	15
β-strand	698-705	8	15
α-helix	760-768	9
α-helix	779-781	3
α-helix	782-785	4
α-helix	791-810	20
α-helix	826-829	4
α-helix	853-855	3
α-helix	858-866	9
α-helix	879-888	10
α-helix	891-900	10
α-helix	904-914	11
α-helix	923-932	10
α-helix	942-945	4
α-helix	949-952	4
α-helix	956-966	11
α-helix	968	1
α-helix	983-993	11
α-helix	996-1013	18
α-helix	1016-1018	3
α-helix	1019-1022	4
α-helix	1026-1039	14
α-helix	1042-1050	9
α-helix	1055-1058	4
α-helix	1061-1076	16
α-helix	1080-1093	14
β-strand	1104-1105	2	16
β-strand	1106	1	17
β-strand	1113	1	17
β-strand	1144	1	18
β-strand	1151	1	18
α-helix	1152-1153	2
β-strand	1154	1	19
α-helix	1155	1
β-strand	1161	1	19
α-helix	1190-1192	3
β-strand	1194-1197	4	20
β-strand	1202	1	21
α-helix	1206-1212	7
α-helix	1229-1231	3

Chain B: 42 helices, 67 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	22
β-strand	17-21	5	22
β-strand	25-32	8	22
α-helix	37-39	3
β-strand	43-44	2	22
β-strand	49-57	9	22
β-strand	63-68	6	23
β-strand	77-82	6	23
β-strand	87-91	5	23
α-helix	101-103	3
β-strand	107-108	2	23
β-strand	116-121	6	24
β-strand	128-133	6	24
β-strand	140-146	7	24
β-strand	179-183	5	24
β-strand	189-192	4	25
β-strand	199-203	5	25
β-strand	207-212	6	25
β-strand	217-223	7	25
β-strand	228-232	5	26
β-strand	239-244	6	26
β-strand	247-252	6	26
β-strand	256	1	27
β-strand	258	1	27
β-strand	261-262	2	26
α-helix	263-266	4
β-strand	270-275	6	28
β-strand	282-287	6	28
β-strand	293-298	6	28
β-strand	313-318	6	28
α-helix	324-325	2
β-strand	327-330	4	29
β-strand	338-341	4	29
β-strand	347-348	2	29
β-strand	357-360	4	30
β-strand	365-370	6	30
β-strand	373-378	6	30
β-strand	388-391	4	30
α-helix	399	1
β-strand	400-405	6	31
β-strand	411-416	6	31
β-strand	421-425	5	31
β-strand	433-438	6	31
β-strand	445-449	5	32
α-helix	453-455	3
β-strand	458-463	6	32
β-strand	468-472	5	32
β-strand	492-494	3	32
β-strand	501-506	6	33
α-helix	509-511	3
β-strand	514-519	6	33
β-strand	523-528	6	33
β-strand	539-545	7	33
β-strand	553-555	3	34
β-strand	566-569	4	34
β-strand	582-588	7	34
β-strand	593-600	8	34
β-strand	609-612	4	35
β-strand	621-626	6	35
β-strand	630-637	8	35
α-helix	648-650	3
β-strand	651-659	9	35
β-strand	666-671	6	36
β-strand	678-682	5	36
β-strand	686-692	7	36
β-strand	698-705	8	36
α-helix	760-769	10
α-helix	779-781	3
α-helix	782-785	4
α-helix	793-806	14
α-helix	807-811	5
α-helix	812	1
α-helix	821-823	3
β-strand	824	1	37
α-helix	826-829	4
α-helix	846-848	3
α-helix	852-855	4
α-helix	860-867	8
α-helix	879-888	10
α-helix	891-900	10
α-helix	904-916	13
α-helix	923-931	9
α-helix	942-949	8
α-helix	950-952	3
α-helix	956-966	11
α-helix	968	1
α-helix	973-976	4
α-helix	983-993	11
α-helix	996-1013	18
α-helix	1016-1018	3
α-helix	1019-1022	4
β-strand	1023	1	37
α-helix	1026-1039	14
α-helix	1044-1047	4
α-helix	1048-1050	3
α-helix	1055-1058	4
α-helix	1061-1076	16
α-helix	1080-1093	14
α-helix	1095-1097	3
β-strand	1103-1107	5	38
β-strand	1113	1	38
α-helix	1150-1153	4
β-strand	1154	1	39
β-strand	1161	1	39
α-helix	1186-1190	5
β-strand	1194	1	40
β-strand	1196-1197	2	41
β-strand	1205	1	40
α-helix	1206-1213	8

Chain C: 18 helices, 68 β-strands

Element	Residues	Length	Sheet
β-strand	22-24	3	42
β-strand	31-34	4	43
β-strand	40-44	5	43
β-strand	49-53	5	43
β-strand	60-64	5	43
α-helix	73-75	3
β-strand	77-82	6	44
β-strand	89-94	6	44
β-strand	99-103	5	44
β-strand	112-116	5	44
β-strand	123-128	6	45
β-strand	136-140	5	45
β-strand	144-146	3	46
β-strand	148	1	45
β-strand	158-160	3	46
β-strand	168-171	4	47
β-strand	179-182	4	47
β-strand	187-189	3	47
β-strand	202-204	3	47
β-strand	210-215	6	48
β-strand	222-227	6	48
β-strand	231-236	6	48
β-strand	243-249	7	48
β-strand	254-259	6	49
α-helix	260	1
β-strand	267-271	5	49
β-strand	278-281	4	49
β-strand	282	1	50
β-strand	285	1	50
β-strand	291-293	3	49
β-strand	300-307	8	42
β-strand	310-318	9	42
β-strand	322-327	6	42
α-helix	328-330	3
β-strand	332	1	49
β-strand	341-345	5	51
β-strand	350-356	7	51
β-strand	395-401	7	51
β-strand	414-416	3	51
β-strand	424-429	6	52
β-strand	435-440	6	52
β-strand	444-450	7	52
β-strand	456-463	8	52
β-strand	469-474	6	53
α-helix	477-479	3
β-strand	482-487	6	53
β-strand	491-498	8	53
β-strand	511-518	8	53
β-strand	525-530	6	54
α-helix	531-532	2
α-helix	533-535	3
β-strand	538-543	6	54
β-strand	547-552	6	54
β-strand	563-569	7	54
β-strand	577-579	3	55
β-strand	590-594	5	55
β-strand	606-612	7	55
β-strand	617-623	7	55
β-strand	633-636	4	56
β-strand	645-650	6	56
β-strand	654-661	8	56
α-helix	662-663	2
α-helix	672-674	3
β-strand	675-683	9	56
β-strand	690-695	6	57
β-strand	702-706	5	57
β-strand	710-716	7	57
β-strand	722-729	8	57
α-helix	781-787	7
α-helix	797-810	14
α-helix	814-827	14
α-helix	1006-1017	12
α-helix	1021-1031	11
α-helix	1032-1034	3
α-helix	1036-1038	3
α-helix	1041-1057	17
α-helix	1061-1070	10
β-strand	1085	1	21
β-strand	1088-1089	2	16
β-strand	1096	1	16
β-strand	1103	1	58
β-strand	1110	1	58
β-strand	1113	1	59
β-strand	1120	1	59
β-strand	1124-1127	4	20
α-helix	1137-1145	9

Chain D: 20 helices, 36 β-strands

Element	Residues	Length	Sheet
α-helix	535-538	4
β-strand	541-545	5	60
β-strand	549-553	5	60
α-helix	566-568	3
β-strand	647-651	5	60
β-strand	653	1	61
β-strand	661-664	4	60
β-strand	672-677	6	62
β-strand	683-688	6	62
β-strand	693-697	5	62
α-helix	699-701	3
β-strand	707-709	3	62
β-strand	716-721	6	63
α-helix	724-726	3
β-strand	729-734	6	63
β-strand	739-745	7	63
β-strand	748-755	8	63
β-strand	762-767	6	64
α-helix	768-769	2
β-strand	775-780	6	64
β-strand	785-792	8	64
β-strand	795-801	7	64
β-strand	809-814	6	65
α-helix	815-817	3
β-strand	837-842	6	65
β-strand	847-852	6	65
β-strand	858-864	7	65
β-strand	871-875	5	66
β-strand	885-891	7	66
β-strand	895-901	7	66
β-strand	913-916	4	66
β-strand	921-925	5	67
β-strand	932-937	6	67
β-strand	942-947	6	67
β-strand	953-955	3	67
β-strand	997	1	61
α-helix	1003-1008	6
α-helix	1016-1030	15
α-helix	1033-1045	13
α-helix	1048-1050	3
α-helix	1062-1064	3
α-helix	1069-1080	12
α-helix	1085-1098	14
α-helix	1179-1191	13
α-helix	1196-1198	3
α-helix	1199-1216	18
α-helix	1220-1226	7
α-helix	1232-1236	5
β-strand	1239-1244	6	38
β-strand	1251-1252	2	38
β-strand	1266	1	68
β-strand	1273	1	68
β-strand	1277-1280	4	41
β-strand	1287-1289	3	41
α-helix	1290-1296	7
α-helix	1311-1315	5

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
GATOR2 complex protein MIOS	A, B	protein	1113	Homo sapiens	Q9NXC5 (AlphaFold model)
GATOR2 complex protein WDR24	C	protein	905	Homo sapiens	Q96S15 (AlphaFold model)
Protein SEC13 homolog	D	protein	633	Homo sapiens	P55735 (AlphaFold model)

Sequence of entity 1 (A, B), FASTA

>9DX0_1 GATOR2 complex protein MIOS (chains A, B)
TKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDTPYM
KCVAWYLNYDPECLLAVGEANGRVVLTSLGQDHNSKFKDLIGKEFVPKHARQCNTLAWNP
LDSDWLAAGLDKHRADFSVLIWDICVTKPLYELGQNDACLSLCWLPRDQKLLLAGMHRNL
AIFDLRNTSQKMFVNTKAVQGVTVDPYFHDRVASFYEGQVAIWDLRKFEKPVLTLTEQPK
PLTKVAWCPTRTGLLATLTRDSNIIRLYDMQHTPTPIGDETEPTIIERSVQPCDNYIASF
AWHPTSQNRMIVVTPNRTMSDFTVFERISLAWSPITSLMWACGRHLYECTEMFVARSIAA
DHKDLIHDVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHSGSVWRVTWAHPE
FGQVLASCSFDRTAAVWEEIVGESNDKLRGQSHWVKRTTLVDSRTSVTDVKFAPKHMGLM
LATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSPMIAVGSDDS
SPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAIATKDVRIFT
LKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDGCVRLWKANY
MDNWKCTGILKGNGSPEKDIATKMRLRALSRYGLDTEQVWRNHILAGNEDPQLKSLWYTL
HFMKQYTEDMDQKSPGNKGSLVYAGIKSIVKSSLGMVESSRHNWSGLDKQSDIQNLNEER
ILALQLCGWIKKGTDVDVGPFLNSLVQEGEWERAAAVALFNLDIRRAIQILNEGASSEKG
DLNLNVVAMALSGYTDEKNSLWREMCSTLRLQLNNPYLCVMFAFLTSETGSYDGVLYENK
VAVRDRVAFACKFLSDTQLNRYIEKLTNEMKEAGNLEGILLTGLTKDGVDLMESYVDRTG
DVQTASYCMLQGSPLDVLKDERVQYWIENYRNLLDAWRFWHKRAEFDIHRSKLDPSSKPL
AQVFVSCNFCGKSISYKVTSCPGCRKPLPRCALCLINMGTPVKDKKLAQFNNWFTWCHNC
RHGGHAGHMLSWFRDHAECPVSACTCKCMQLDT

Sequence of entity 2 (C), FASTA

>9DX0_2 GATOR2 complex protein WDR24 (chains C)
SVLTGRTMHCHLDAPANAISVCRDAAQVVVAGRSIFKIYAIEEEQFVEKLNLRVGRKPSL
NLSCADVVWHQMDENLLATAATNGVVVTWNLGRPSRNKQDQLFTEHKRTVNKVCFHPTEA
HVLLSGSQDGFMKCFDLRRKDSVSTFSGQSESVRDVQFSIRDYFTFASTFENGNVQLWDI
RRPDRCERMFTAHNGPVFCCDWHPEDRGWLATGGRDKMVKVWDMTTHRAKEMHCVQTIAS
VARVKWRPECRHHLATCSMMVDHNIYVWDVRRPFVPAAMFEEHRDVTTGIAWRHPHDPSF
LLSGSKDSSLCQHLFRDASQPVERANPEGLCYGLFGDLAFAAKESLVFAGLASSALSVFE
TEPMFVARSIAADHKDLIHDVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHS
GSVWRVTWAHPEFGQVLASCSFDRTAAVWEEIVGQSHWVKRTTLVDSRTSVTDVKFAPKH
MGLMLATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSPMIAVG
SDDSSPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAIATKDV
RIFTLKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDGCVRLW
KANYMDNWKCTGILKGGMRWFVDTAERYALAGRPLAELCDHNAKVARELGRNQVAQTWTM
LRIIYCSPFSLLSVSHALYDSRLPPDFFGVLVRDMLHFYAEQGDVQMAVSVLIVLGERVR
KDIDEQTQEHWYTSYIDLLQRFRLWNVSNEVVKLSTSRAVSCLNQASTTLHVNCSHCKRP
MSSRGWVCDRCHRCASMCAVCHHVVKGLFVWCQGCSHGGHLQHIMKWLEGSSHCPAGCGH
LCEYS

Sequence of entity 3 (D), FASTA

>9DX0_3 Protein SEC13 homolog (chains D)
ARVTTAYGSYQDANIPFPRTSGARFCGAGYLVYFTRPMTMHRAVSPTEPTPRSLSALSAY
HTSKREGSDSGNRQIKAAGKVIIQDGSGSGMVSVINTVDTSHEDMIHDAQMDYYGTRLAT
CSSDRSVKIFDVRNGGQILIADLRGHEGPVWQVAWAHPMYGNILASCSYDRKVIIWREEN
GTWEKSHEHAGHDSSVNSVCWAPHDYGLILACGSSDGAISLLTYTGEGQWEVKKINNAHT
IGCNAVSWAPAVVPDHPSGQKPNYIKRFASGGCDNLIKLWKEEEDGQWKEEQKLEAHSDW
VRDVAWAPSIGLPTSTIASCSQDGRVFIWTCDDASSNTWSPKLLHKFNDVVWHVSWSITA
NILAVSGGDNKVTLWKESVDGQWVCISDIACLLPVHKSLGELYILNVNDIQETCQKNAAS
ALLVGRKDLVQVWSLATVATDLCLGPKSDPDLETPWARHPFGRQLLESLLAHYCRLRDVQ
TLAMLCSVFEAQSDPRERERDQHDKNKRLLDPANTQQFDDFKKCYGEILYRWGLREKRAE
VLKFVSCPPDPHKGIEFGVYCSHCRSEVRGTQCAICKGFTFQCAICHVAVRGSSNFCLTC
GHGGHTSHMMEWFRTQEVCPTGCGCHCLLESTF

Primary citation

Structural basis for the dynamic regulation of mTORC1 by amino acids. Valenstein, M.L., Wranik, M., Lalgudi, P.V. et al. Nature (2025) 646:493-500. DOI 10.1038/s41586-025-09428-7 · 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
9OTI 3.5 Å, GATOR2 complex bound to arginine sensor CASTOR1
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

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