9DX2: Human GATOR2 complex - CASTOR1 bound state

Human GATOR2 complex - CASTOR1 bound state. Determined by electron microscopy at 3.4 Å resolution. Released 20 May 2026.

Method: Electron microscopy
Resolution: 3.4 Å
Organism: Homo sapiens
Chains: 8
Atoms: 34,252
Mol. weight: 529.19 kDa
Released: 20 May 2026

Explore 9DX2 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9DX2 contains 146 α-helices and 298 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.

Chain A: 33 helices, 38 β-strands

Element	Residues	Length	Sheet
β-strand	356-360	5	1
β-strand	366-370	5	1
β-strand	374-377	4	1
β-strand	389-391	3	1
β-strand	400-404	5	2
β-strand	411-416	6	2
β-strand	420-426	7	2
β-strand	432-439	8	2
β-strand	445-450	6	3
α-helix	453-455	3
β-strand	459-463	5	3
β-strand	468-474	7	3
β-strand	487-494	8	3
β-strand	501-506	6	4
α-helix	507-508	2
α-helix	509-511	3
β-strand	514-519	6	4
β-strand	523-528	6	4
β-strand	539-545	7	4
β-strand	550-555	6	5
β-strand	566-571	6	5
β-strand	582-588	7	5
β-strand	593-599	7	5
β-strand	609-612	4	6
β-strand	621-626	6	6
β-strand	630-637	8	6
α-helix	648-650	3
β-strand	651-659	9	6
β-strand	666-671	6	7
β-strand	678-682	5	7
β-strand	686-692	7	7
β-strand	698-705	8	7
α-helix	760-768	9
α-helix	779-781	3
α-helix	782-786	5
α-helix	791-812	22
α-helix	822-824	3
α-helix	826-829	4
α-helix	830-832	3
α-helix	853-855	3
α-helix	858-867	10
α-helix	879-886	8
α-helix	891-900	10
α-helix	904-915	12
α-helix	922-932	11
α-helix	941-949	9
α-helix	950-952	3
α-helix	956-966	11
α-helix	973-976	4
α-helix	983-993	11
α-helix	996-1011	16
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	8
β-strand	1106	1	9
β-strand	1113	1	9
β-strand	1144	1	10
β-strand	1151	1	10
α-helix	1152-1153	2
β-strand	1154	1	11
β-strand	1161	1	11
α-helix	1190-1192	3
β-strand	1194-1197	4	12
β-strand	1202	1	13
β-strand	1203-1205	3	12
α-helix	1206-1213	8

Chain B: 29 helices, 41 β-strands

Element	Residues	Length	Sheet
β-strand	357-361	5	14
β-strand	365-369	5	14
β-strand	374-377	4	14
β-strand	400-405	6	15
β-strand	411-416	6	15
β-strand	420-425	6	15
β-strand	433-439	7	15
β-strand	445-450	6	16
α-helix	453-455	3
β-strand	458	1	17
β-strand	459-463	5	16
β-strand	467-469	3	16
β-strand	472	1	17
β-strand	489	1	17
β-strand	501-506	6	18
α-helix	507-508	2
α-helix	509-511	3
β-strand	514-519	6	18
β-strand	524-528	5	18
β-strand	542-544	3	18
β-strand	552-555	4	19
β-strand	566-570	5	19
β-strand	582-588	7	19
β-strand	593-599	7	19
β-strand	609-612	4	20
β-strand	621-626	6	20
β-strand	630-635	6	20
β-strand	641	1	21
β-strand	643	1	21
β-strand	655-659	5	20
β-strand	668-671	4	22
β-strand	678-681	4	22
β-strand	686	1	23
β-strand	687-692	6	22
β-strand	698-700	3	22
β-strand	705	1	23
α-helix	760-769	10
α-helix	779-781	3
α-helix	782-785	4
α-helix	791-812	22
α-helix	820-823	4
β-strand	824	1	24
α-helix	826-829	4
α-helix	858-867	10
α-helix	880-888	9
α-helix	891-900	10
α-helix	905-915	11
α-helix	924-931	8
α-helix	941-949	9
α-helix	956-965	10
α-helix	973-976	4
α-helix	983-990	8
α-helix	996-1010	15
α-helix	1016-1018	3
α-helix	1019-1022	4
β-strand	1023	1	24
α-helix	1028-1039	12
α-helix	1043-1049	7
α-helix	1055-1058	4
α-helix	1061-1076	16
α-helix	1080-1093	14
β-strand	1103-1104	2	25
α-helix	1150-1153	4
β-strand	1154	1	26
β-strand	1161	1	26
α-helix	1186-1190	5
β-strand	1194	1	27
β-strand	1196-1197	2	28
β-strand	1205	1	27
α-helix	1206-1215	10

Chain C: 15 helices, 35 β-strands

Element	Residues	Length	Sheet
β-strand	355-356	2	29
β-strand	395-396	2	29
β-strand	397-401	5	30
β-strand	412-416	5	30
β-strand	424-429	6	31
β-strand	435-440	6	31
β-strand	444-450	7	31
β-strand	456-463	8	31
β-strand	469-474	6	32
α-helix	477-479	3
β-strand	482-487	6	32
β-strand	492-498	7	32
β-strand	511-518	8	32
β-strand	525-530	6	33
α-helix	533-535	3
β-strand	538-543	6	33
β-strand	547-552	6	33
β-strand	563-569	7	33
β-strand	577-579	3	34
β-strand	590-594	5	34
β-strand	606-612	7	34
β-strand	617-623	7	34
β-strand	633-636	4	35
β-strand	645-650	6	35
β-strand	654-661	8	35
α-helix	662-663	2
α-helix	672-674	3
β-strand	675-683	9	35
β-strand	690-695	6	36
β-strand	701-706	6	36
β-strand	710-716	7	36
β-strand	722-729	8	36
α-helix	781-789	9
α-helix	797-811	15
α-helix	814-827	14
α-helix	999-1000	2
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	1061-1069	9
β-strand	1085	1	13
β-strand	1088-1089	2	8
β-strand	1096	1	8
β-strand	1113	1	37
β-strand	1120	1	37
β-strand	1124-1127	4	12
β-strand	1136	1	12
α-helix	1137-1146	10

Chain D: 20 helices, 28 β-strands

Element	Residues	Length	Sheet
α-helix	535-538	4
β-strand	541-545	5	38
β-strand	549-553	5	38
α-helix	566-568	3
α-helix	631-634	4
β-strand	647-651	5	38
β-strand	661-664	4	38
β-strand	672-677	6	39
β-strand	683-688	6	39
β-strand	693-697	5	39
α-helix	699-701	3
β-strand	704-709	6	39
β-strand	716-721	6	40
α-helix	724-726	3
β-strand	729-734	6	40
β-strand	739-744	6	40
β-strand	749-755	7	40
β-strand	762-767	6	41
α-helix	768-769	2
β-strand	775-780	6	41
β-strand	785-792	8	41
β-strand	795-801	7	41
β-strand	809-814	6	42
α-helix	815-816	2
α-helix	827-829	3
β-strand	837-842	6	42
β-strand	847-852	6	42
β-strand	858-864	7	42
β-strand	871-876	6	43
β-strand	885-891	7	43
β-strand	895-901	7	43
β-strand	910-916	7	43
β-strand	921-925	5	44
β-strand	933-937	5	44
β-strand	942-947	6	44
β-strand	953-955	3	44
α-helix	1003-1006	4
α-helix	1016-1030	15
α-helix	1033-1045	13
α-helix	1051-1053	3
α-helix	1062-1064	3
α-helix	1069-1082	14
α-helix	1085-1098	14
α-helix	1179-1190	12
α-helix	1196-1198	3
α-helix	1199-1215	17
α-helix	1220-1225	6
α-helix	1226-1228	3

Chain E: 14 helices, 19 β-strands

Element	Residues	Length	Sheet
β-strand	2-15	14	45
α-helix	20-23	4
α-helix	24-29	6
α-helix	34-36	3
β-strand	41-46	6	45
β-strand	51-56	6	45
α-helix	59-62	4
β-strand	69-71	3	45
β-strand	76-80	5	45
α-helix	96-97	2
α-helix	98-102	5
α-helix	103-107	5
β-strand	113-116	4	45
β-strand	121-126	6	45
α-helix	130-137	8
β-strand	142-146	5	45
β-strand	151-153	3	45
α-helix	169-171	3
β-strand	177	1	45
β-strand	184-187	4	45
β-strand	188	1	46
α-helix	191-197	7
α-helix	198-205	8
β-strand	228-233	6	45
β-strand	236-242	7	45
α-helix	245-248	4
β-strand	255	1	46
β-strand	257	1	47
β-strand	263-268	6	45
β-strand	296-299	4	45
β-strand	304-309	6	45
α-helix	310-312	3
α-helix	313-326	14

Chain F: 14 helices, 20 β-strands

Element	Residues	Length	Sheet
β-strand	2-5	4	48
β-strand	10-15	6	48
α-helix	20-23	4
α-helix	24-31	8
β-strand	41-46	6	48
β-strand	51-56	6	48
α-helix	58-62	5
β-strand	69-71	3	48
β-strand	76-80	5	48
α-helix	98-102	5
α-helix	103-107	5
β-strand	112-116	5	48
β-strand	121-126	6	48
α-helix	127-129	3
α-helix	130-137	8
β-strand	142-146	5	48
β-strand	151-153	3	48
β-strand	178	1	48
β-strand	184-187	4	48
β-strand	188	1	49
α-helix	191-193	3
α-helix	195-197	3
α-helix	198-205	8
β-strand	228-233	6	48
β-strand	236-242	7	48
α-helix	243-246	4
β-strand	255	1	49
β-strand	257	1	50
β-strand	263-268	6	48
α-helix	282-290	9
β-strand	296-299	4	48
β-strand	304-309	6	48
α-helix	310-312	3
α-helix	313-326	14

Chain G: 13 helices, 18 β-strands

Element	Residues	Length	Sheet
β-strand	3-4	2	51
β-strand	10-15	6	52
α-helix	21-23	3
α-helix	24-31	8
α-helix	34-36	3
β-strand	41-46	6	52
β-strand	51-55	5	52
α-helix	59-61	3
β-strand	69-71	3	52
β-strand	76-79	4	52
α-helix	96-99	4
α-helix	102-108	7
β-strand	113-116	4	52
β-strand	121-126	6	52
α-helix	130-137	8
β-strand	143-144	2	51
β-strand	145-146	2	53
β-strand	151-152	2	53
β-strand	177-178	2	52
β-strand	184-187	4	52
α-helix	191-197	7
α-helix	198-205	8
β-strand	228-233	6	52
β-strand	236-242	7	52
α-helix	243-246	4
β-strand	263-268	6	52
α-helix	280-291	12
β-strand	296-299	4	52
β-strand	304-309	6	52
α-helix	310-312	3
α-helix	313-326	14

Chain H: 8 helices, 99 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	54
β-strand	17-21	5	54
β-strand	25-32	8	54
β-strand	43-44	2	54
β-strand	49-57	9	54
β-strand	63-68	6	55
β-strand	77-82	6	55
β-strand	87-91	5	55
β-strand	108	1	55
β-strand	112	1	50
β-strand	116-121	6	56
β-strand	128-133	6	56
β-strand	140-146	7	56
β-strand	150	1	55
β-strand	155-159	5	56
β-strand	165-168	4	57
α-helix	169	1
β-strand	175-179	5	57
β-strand	183	1	58
β-strand	184-188	5	57
β-strand	193-194	2	57
β-strand	199	1	58
β-strand	204-208	5	59
β-strand	215-219	5	59
β-strand	224-228	5	59
β-strand	237-238	2	59
α-helix	239-242	4
β-strand	246-251	6	60
β-strand	258-263	6	60
β-strand	269-273	5	60
α-helix	284-285	2
β-strand	286-290	5	60
β-strand	299-302	4	61
β-strand	309-313	5	61
β-strand	319-323	5	61
β-strand	330-332	3	62
β-strand	339-342	4	62
β-strand	347-354	8	62
β-strand	365-368	4	62
β-strand	371-379	9	62
β-strand	385-390	6	63
β-strand	399-404	6	63
β-strand	409-413	5	63
α-helix	423-425	3
β-strand	429-430	2	63
β-strand	434	1	47
β-strand	438-443	6	64
β-strand	450-455	6	64
β-strand	462-468	7	64
β-strand	472	1	63
β-strand	477-481	5	64
β-strand	489-490	2	65
β-strand	497-501	5	65
β-strand	505-510	6	65
β-strand	515-521	7	65
β-strand	526-530	5	66
β-strand	537-542	6	66
β-strand	545-550	6	66
β-strand	559-560	2	66
β-strand	568-573	6	67
β-strand	580-585	6	67
β-strand	592-595	4	67
β-strand	602-604	3	67
β-strand	614-617	4	68
β-strand	624-629	6	68
β-strand	633-638	6	68
β-strand	647-650	4	69
β-strand	655-660	6	70
β-strand	666-671	6	70
β-strand	675-680	6	70
β-strand	685-690	6	70
α-helix	699-701	3
β-strand	703-708	6	71
β-strand	716-720	5	71
β-strand	725-728	4	71
β-strand	738	1	71
β-strand	749-754	6	72
β-strand	761-766	6	72
β-strand	770-775	6	72
β-strand	783-786	4	72
β-strand	792-797	6	73
β-strand	804-809	6	73
β-strand	813-818	6	73
β-strand	825-830	6	73
β-strand	836-841	6	74
β-strand	848-853	6	74
β-strand	857-862	6	74
β-strand	869-875	7	74
β-strand	880-885	6	75
β-strand	892-897	6	75
β-strand	904-908	5	75
β-strand	911	1	75
β-strand	917-919	3	75
β-strand	926-931	6	69
β-strand	939-944	6	69
β-strand	948-953	6	69
α-helix	954-956	3
β-strand	958	1	75
β-strand	970-971	2	30
β-strand	976-978	3	30
β-strand	983-984	2	25
β-strand	994	1	76
β-strand	1001	1	76
β-strand	1005-1008	4	28
β-strand	1015-1017	3	28
α-helix	1018-1027	10
α-helix	1039-1043	5

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
GATOR2 complex protein MIOS	A, B	protein	786	Homo sapiens	Q9NXC5 (AlphaFold model)
Nucleoporin SEH1	C	protein	570	Homo sapiens	Q96EE3 (AlphaFold model)
Protein SEC13 homolog	D	protein	545	Homo sapiens	P55735 (AlphaFold model)
Cytosolic arginine sensor for mTORC1 subunit 1	E, F, G	protein	328	Homo sapiens	Q8WTX7 (AlphaFold model)
GATOR2 complex protein WDR24	H	protein	1042	Homo sapiens	Q96S15

Sequence of entity 1 (A, B), FASTA

>9DX2_1 GATOR2 complex protein MIOS (chains A, B)
ERISLAWSPITSLMWACGRHLYECTEMFVARSIAADHKDLIHDVSFDFHGRRMATCSSDQ
SVKVWDKSESGDWHCTASWKTHSGSVWRVTWAHPEFGQVLASCSFDRTAAVWEEIVGESN
DKLRGQSHWVKRTTLVDSRTSVTDVKFAPKHMGLMLATCSADGIVRIYEAPDVMNLSQWS
LQHEISCKLSCSCISWNPSSSRAHSPMIAVGSDDSSPNAMAKVQIFEYNENTRKYAKAET
LMTVTDPVHDIAFAPNLGRSFHILAIATKDVRIFTLKPVRKELTSSGGPTKFEIHIVAQF
DNHNSQVWRVSWNITGTVLASSGDDGCVRLWKANYMDNWKCTGILKGNGSPVEKDIATKM
RLRALSRYGLDTEQVWRNHILAGNEDPQLKSLWYTLHFMKQYTEDMDQKSPGNKGSLVYA
GIKSIVKSSLGMVESSRHNWSGDIQNLNEERILALQLCGWIKKGTDVDVGPFLNSLVQEG
EWERAAAVALFNLDIRRAIQILNEGASSEKGDLNLNVVAMALSGYTDEKNSLWREMCSTL
RLQLNNPYLCVMFAFLTSETGSYDGVLYENKVAVRDRVAFACKFLSDTQLNRYIEKLTNE
MKEAGNLEGILLTGLTKDGVDLMESYVDRTGDVQTASYCMLQGSPLDVLKDERVQYWIEN
YRNLLDAWRFWHKRAEFDIHRSKLDPSSKPLAQVFVSCNFCGKSISYSKVTSCPGCRKPL
PRCALCLINMGTPVSKDKKLAQFNNWFTWCHNCRHGGHAGHMLSWFRDHAECPVSACTCK
CMQLDT

Sequence of entity 2 (C), FASTA

>9DX2_2 Nucleoporin SEH1 (chains C)
AAKESLVAEPFAGLASSALSVFETEPMFVARSIAADHKDLIHDVSFDFHGRRMATCSSDQ
SVKVWDKSESGDWHCTASWKTHSGSVWRVTWAHPEFGQVLASCSFDRTAAVWEEIVSHWV
KRTTLVDSRTSVTDVKFAPKHMGLMLATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLS
CSCISWNPSSSRAHSPMIAVGSDDSSPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHD
IAFAPNLGRSFHILAIATKDVRIFTLKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRV
SWNITGTVLASSGDDGCVRLWKANYMDNWKCTGILKGGGMRWFVDTAERYALAGRPLAEL
CDHNAKVARELGRNQVAQTWTMLRIIYCSPSSSFSLLSVSHALYDSRLPPDFFGVLVRDM
LHFYAEQGDVQMAVSVLIVLGERVRKDIDEQTQEHWYTSYIDLLQRFRLWNVSNEVVKLS
TSRAVSCLNQASTTLHVNCSHCKRPMSSRGWVCDRCHRCASMCAVCHHVVKGLFVWCQGC
SHGGHLQHIMKWLEGSSHCPAGCGHLCEYS

Sequence of entity 3 (D), FASTA

>9DX2_3 Protein SEC13 homolog (chains D)
AYGSYQDANIPFPRTSGARFCGAGYLVYFTRPMTMHRAVSPTEPTPRSLSALSAYHTKRE
GSDSGNRQIKAAGKVIIQDGSGSGMVSVINTVDTSHEDMIHDAQMDYYGTRLATCSSDRS
VKIFDVRNGGQILIADLRGHEGPVWQVAWAHPMYGNILASCSYDRKVIIWREENGTWEKS
HEHAGHDSSVNSVCWAPHDYGLILACGSSDGAISLLTYTGEGQWEVKKINNAHTIGCNAV
SWAPAVVPGSLIDHPSGQKPNYIKRFASGGCDNLIKLWKEEEDGQWKEEQKLEAHSDWVR
DVAWAPSIGLPTSTIASCSQDGRVFIWTCDDASSNTWSPKLLHKFNDVVWHVSWSITANI
LAVSGGDNKVTLWKESVDGQWVCISIACLLPVHKSLGELYILNVNDIQETCQKNAASALL
VGRKDLVQVWSLATVATDLCLGPKSDPDLETPWARHPFGRQLLESLLAHYCRLRDVQTLA
MLCSVFEAQSLDPRERERDQHDKNKRLLDPANTQQFDDFKKCYGEILYRWGLREKRAEVL
KFVSC

Sequence of entity 4 (E, F, G), FASTA

>9DX2_4 Cytosolic arginine sensor for mTORC1 subunit 1 (chains E, F, G)
MELHILEHRVRVLSVARPGLWLYTHPLIKLLFLPRRSRCKFFSLTETPEDYTLMVDEEGF
KELPPSEFLQVAEATWLVLNVSSHSGAAVQAAGVTKIARSVIAPLAEHHVSVLMLSTYQT
DFILVREQDLSVVIHTLAQEFDIYREVGGEPVPVTRDDSSNGFPRTQHGPSPTVHPIQSP
QNRFCVLTLDPETLPAIATTLIDVLFYSHSTPKEAASSSPEPSSITFFAFSLIEGYISIV
MDAETQKKFPSDLLLTSSSGELWRMVRIGGQPLGFDECGIVAQIAGPLAAADISAYYIST
FNFDHALVPEDGIGSVIEVLQRRQEGLA

Sequence of entity 5 (H), FASTA

>9DX2_5 GATOR2 complex protein WDR24 (chains H)
TKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDTPYM
KCVAWYLNYDPECLLAVGEANGRVVLTSLGQDHNSKFKDLIGKEYVPKHARQCNTLAWNP
LDSDWLAAGLDKHRADFSVLIWDICLLTKPLYELGQNDACLSLCWLPRDQKLLLAGMHRN
LAIFDLRNTSEKMFVNTKAVQGVTVDPYFHDRVASFYEGQVAIWDLRKFEKPVLTLTEQP
KPLTKVAWCPTRTGLLATLTRDSNIIRLYDMQHTPTPIGPTIIERSVQPCDNYIASFAWH
PTSQNRMIVVTPNRTMSDFTVFTKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAG
SLRLSEDSAATLLSINSDTPYMKCVAWYLNYDPECLLAVGQANGRVVLTSLGQDHNSKFK
DLIGKEFVPKHARQCNTLAWNPLDSNWLAAGLDKHRADFSVLIWDICLVTKPLYELGQND
ACLSLCWLPRDQKLLLAGMHRNLAIFDLRNTSQKMFVNTKAVQGVTVDPYFHDRVASFYE
GQVAIWDLRKFEKPVLTLTEQPKPLTKVAWCPTRTGLLATLTRDSNIIRLYDMQHTIIER
SVQPCDDYIASFAWHPTSQNRMIVVTPNRTMSDFTVFVLTGRTMHCHLDAPANAISVCRD
AAQVVVAGRSIFKIYAIEEEQFVEKLNLRVGRKPSLNLSCADVVWHQMDENLLATAATNG
VVVTWNLGRPSRNKQDQLFTEHKRTVNKVCFHPTEAHVLLSGSQDGFMKCFDLRRKDSVS
TFSGQSESVRDVQFSIRDYFTFASTFENGNVQLWDIRRPDRCERMFTAHNGPVFCCDWHP
EDRGWLATGGRDKMVKVWDMTTHRAKEMHCVQTIASVARVKWRPECRHHLATCSMMVDHN
IYVWDVRRPFVPAAMFEEHRDVTTGIAWRHPHDPSFLLSGSKDSSLCQHLFRDASQPVER
ANPEGLCYGLFGDLAFPPFGVYCSHCRSTFQCAICHVAVRGSSNFCLTCGHGGHTSHMME
WFRTQEVCPTGCGCHCLLESTF

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
9LWF 3.41 Å, Cryo-EM structure of dual sensor bound GATOR2 complex
9DX0 3.47 Å, Human GATOR2 complex - apo state
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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