9DX1: Human GATOR2 complex - Sestrin2 bound state

Human GATOR2 complex - Sestrin2 bound state. Determined by electron microscopy at 3.36 Å resolution. Released 20 May 2026.

Method: Electron microscopy
Resolution: 3.36 Å
Organism: Homo sapiens
Chains: 6
Atoms: 30,499
Mol. weight: 503.76 kDa
Released: 20 May 2026

Explore 9DX1 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9DX1 contains 133 α-helices and 241 β-strands across 6 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, 36 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	400-405	6	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	458-463	6	3
β-strand	468-475	8	3
α-helix	476-477	2
β-strand	486-494	9	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	582-587	6	5
β-strand	594-600	7	5
β-strand	609-612	4	6
β-strand	621-626	6	6
β-strand	630-637	8	6
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	760-769	10
α-helix	779-781	3
α-helix	782-786	5
α-helix	793-811	19
α-helix	826-829	4
α-helix	830-832	3
α-helix	853-855	3
α-helix	858-867	10
α-helix	879-888	10
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	996-1013	18
α-helix	1016-1018	3
α-helix	1019-1022	4
α-helix	1026-1039	14
α-helix	1042-1050	9
α-helix	1057-1059	3
α-helix	1061-1076	16
α-helix	1080-1093	14
β-strand	1104-1106	3	8
β-strand	1113	1	8
β-strand	1144	1	9
β-strand	1151	1	9
α-helix	1152-1153	2
β-strand	1154	1	10
α-helix	1155	1
β-strand	1161	1	10
α-helix	1190-1192	3
β-strand	1194-1197	4	11
β-strand	1202	1	12
β-strand	1203-1205	3	11
α-helix	1206-1213	8

Chain B: 35 helices, 69 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	13
β-strand	17-21	5	13
β-strand	25-32	8	13
α-helix	37-39	3
β-strand	43-44	2	13
β-strand	49-57	9	13
β-strand	66-68	3	14
β-strand	77-81	5	14
β-strand	87-91	5	14
α-helix	101-103	3
β-strand	107-108	2	14
β-strand	116-121	6	15
β-strand	128-133	6	15
β-strand	140-146	7	15
β-strand	179-183	5	15
β-strand	189-192	4	16
β-strand	199-203	5	16
β-strand	207-212	6	16
β-strand	217-223	7	16
β-strand	228-232	5	17
β-strand	239-244	6	17
β-strand	247-252	6	17
β-strand	256	1	18
β-strand	258	1	18
β-strand	261-262	2	17
α-helix	263-266	4
β-strand	270-275	6	19
β-strand	282-287	6	19
β-strand	293-298	6	19
β-strand	313-318	6	19
α-helix	324-325	2
β-strand	327-330	4	20
β-strand	338-341	4	20
β-strand	347-348	2	20
β-strand	357-360	4	21
β-strand	366-370	5	21
β-strand	373-377	5	21
β-strand	391	1	21
α-helix	399	1
β-strand	400-404	5	22
β-strand	411-416	6	22
β-strand	420-425	6	22
β-strand	433-439	7	22
β-strand	445-450	6	23
β-strand	458-463	6	23
β-strand	468-472	5	23
β-strand	489-494	6	23
β-strand	503-506	4	24
α-helix	507-508	2
α-helix	509-511	3
β-strand	514-518	5	24
β-strand	524-528	5	24
β-strand	539-543	5	24
β-strand	550-555	6	25
β-strand	566-571	6	25
β-strand	582-588	7	25
β-strand	593-599	7	25
β-strand	609-612	4	26
β-strand	621-626	6	26
β-strand	630-637	8	26
α-helix	638-639	2
β-strand	651-659	9	26
β-strand	666-671	6	27
β-strand	678-682	5	27
β-strand	686-692	7	27
β-strand	698-705	8	27
α-helix	760-769	10
α-helix	779-783	5
α-helix	791-812	22
β-strand	824	1	28
α-helix	826-829	4
α-helix	851-855	5
α-helix	858-867	10
α-helix	879-888	10
α-helix	891-900	10
α-helix	904-913	10
α-helix	923-931	9
α-helix	941-949	9
α-helix	950-952	3
α-helix	956-966	11
α-helix	983-993	11
α-helix	996-1013	18
α-helix	1016-1018	3
α-helix	1019-1022	4
β-strand	1023	1	28
α-helix	1026-1039	14
α-helix	1043-1049	7
α-helix	1055-1058	4
α-helix	1061-1076	16
α-helix	1080-1091	12
α-helix	1095-1097	3
β-strand	1103-1107	5	29
β-strand	1113	1	29
β-strand	1144	1	30
α-helix	1150	1
β-strand	1151	1	30
α-helix	1152-1153	2
β-strand	1154	1	31
β-strand	1161	1	31
α-helix	1186-1190	5
β-strand	1194	1	32
β-strand	1196-1197	2	33
β-strand	1205	1	32
α-helix	1206-1212	7

Chain C: 14 helices, 34 β-strands

Element	Residues	Length	Sheet
β-strand	342-345	4	34
β-strand	350-356	7	34
β-strand	395-401	7	34
β-strand	412-416	5	34
β-strand	424-429	6	35
β-strand	435-440	6	35
β-strand	445-450	6	35
β-strand	456-462	7	35
β-strand	469-474	6	36
β-strand	482-487	6	36
β-strand	492-498	7	36
β-strand	511-518	8	36
β-strand	525-530	6	37
α-helix	531-532	2
α-helix	533-535	3
β-strand	538-543	6	37
β-strand	547-552	6	37
β-strand	563-569	7	37
β-strand	576-579	4	38
β-strand	590-594	5	38
β-strand	606-612	7	38
β-strand	617-624	8	38
β-strand	633-636	4	39
β-strand	645-650	6	39
β-strand	654-661	8	39
α-helix	662-663	2
α-helix	672-674	3
β-strand	675-683	9	39
β-strand	690-695	6	40
β-strand	702-706	5	40
β-strand	710-716	7	40
β-strand	722-729	8	40
α-helix	781-789	9
α-helix	797-811	15
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	1061-1069	9
β-strand	1085	1	12
β-strand	1087-1090	4	8
β-strand	1095-1096	2	8
β-strand	1113	1	41
β-strand	1120	1	41
β-strand	1124-1127	4	11
α-helix	1137-1144	8

Chain D: 17 helices, 31 β-strands

Element	Residues	Length	Sheet
α-helix	535-538	4
β-strand	541-545	5	42
β-strand	549-553	5	42
β-strand	646-651	6	42
β-strand	661-665	5	42
β-strand	672-677	6	43
β-strand	683-688	6	43
β-strand	693-697	5	43
α-helix	699-701	3
β-strand	704-709	6	43
β-strand	716-721	6	44
α-helix	724-726	3
β-strand	729-734	6	44
β-strand	739-745	7	44
β-strand	748-755	8	44
β-strand	762-767	6	45
β-strand	775-780	6	45
β-strand	785-790	6	45
β-strand	796-801	6	45
β-strand	809-814	6	46
α-helix	815-817	3
α-helix	819-821	3
β-strand	826	1	47
β-strand	829	1	47
β-strand	837-842	6	46
β-strand	847-852	6	46
β-strand	858-864	7	46
β-strand	871-875	5	48
β-strand	886-891	6	48
β-strand	895-900	6	48
β-strand	910	1	48
β-strand	913-916	4	48
β-strand	921-925	5	49
β-strand	933-937	5	49
β-strand	942-947	6	49
β-strand	953-955	3	49
α-helix	1003-1006	4
α-helix	1016-1030	15
α-helix	1033-1045	13
α-helix	1048-1050	3
α-helix	1062-1064	3
α-helix	1066-1070	5
α-helix	1071-1081	11
α-helix	1085-1099	15
α-helix	1179-1191	13
α-helix	1196-1198	3
α-helix	1199-1215	17
α-helix	1219-1226	8

Chain E: 25 helices, 4 β-strands

Element	Residues	Length	Sheet
β-strand	45	1	50
α-helix	50-54	5
α-helix	78-83	6
α-helix	87-102	16
α-helix	109-123	15
α-helix	126-137	12
α-helix	143-147	5
α-helix	149-151	3
α-helix	154-157	4
α-helix	160-168	9
α-helix	170-172	3
α-helix	175-181	7
α-helix	191-214	24
β-strand	216	1	50
α-helix	217-218	2
α-helix	312-315	4
α-helix	335-336	2
β-strand	337-338	2	51
α-helix	339-341	3
α-helix	344-348	5
α-helix	349-355	7
α-helix	357-370	14
α-helix	386-401	16
β-strand	403-404	2	51
α-helix	412-416	5
α-helix	419-430	12
α-helix	432-434	3
α-helix	437-442	6
α-helix	449-479	31

Chain F: 9 helices, 67 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	52
β-strand	18-21	4	52
β-strand	25-32	8	52
α-helix	37-39	3
β-strand	42-46	5	52
β-strand	49-57	9	52
β-strand	65-68	4	53
β-strand	78-81	4	53
β-strand	87-90	4	53
β-strand	108	1	53
β-strand	116-121	6	54
β-strand	128-133	6	54
β-strand	140-146	7	54
β-strand	152-154	3	54
β-strand	158-163	6	55
β-strand	170-175	6	55
β-strand	178	1	56
β-strand	179-183	5	55
β-strand	194	1	56
β-strand	199-203	5	57
β-strand	210-215	6	57
β-strand	218-223	6	57
β-strand	232-233	2	57
α-helix	234-236	3
β-strand	241-246	6	58
β-strand	253-258	6	58
β-strand	264-269	6	58
β-strand	284-289	6	58
α-helix	295-296	2
β-strand	298-301	4	59
β-strand	308-312	5	59
β-strand	318-322	5	59
β-strand	332-335	4	60
β-strand	342-345	4	61
β-strand	351-355	5	61
β-strand	360-365	6	61
β-strand	370-375	6	61
α-helix	384-386	3
β-strand	388-393	6	62
β-strand	400-405	6	62
β-strand	410-414	5	62
β-strand	423-427	5	62
β-strand	434-438	5	63
β-strand	446-451	6	63
β-strand	455-460	6	63
β-strand	466-471	6	63
β-strand	477-482	6	64
β-strand	489-494	6	64
β-strand	498-503	6	64
β-strand	512-515	4	64
β-strand	521-526	6	65
β-strand	533-538	6	65
β-strand	542-547	6	65
β-strand	554-560	7	65
β-strand	565-570	6	66
α-helix	571	1
β-strand	577-582	6	66
β-strand	589-593	5	66
β-strand	596	1	66
β-strand	602-604	3	66
β-strand	611-618	8	60
β-strand	621-629	9	60
β-strand	633-638	6	60
α-helix	639-641	3
β-strand	643	1	66
α-helix	648-651	4
β-strand	654-658	5	29
β-strand	667	1	29
β-strand	671-672	2	67
β-strand	676-678	3	67
β-strand	681	1	68
β-strand	688	1	68
β-strand	692-695	4	33
β-strand	702-704	3	33
α-helix	705-711	7
α-helix	726-730	5

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
GATOR2 complex protein MIOS	A, B	protein	1116	Homo sapiens	Q9NXC5 (AlphaFold model)
Nucleoporin SEH1	C	protein	595	Homo sapiens	Q96EE3 (AlphaFold model)
Protein SEC13 homolog	D	protein	564	Homo sapiens	P55735 (AlphaFold model)
Sestrin2	E	protein	335	Homo sapiens	P58004 (AlphaFold model)
GATOR2 complex protein WDR24	F	protein	729	Homo sapiens	Q96S15

Sequence of entity 1 (A, B), FASTA

>9DX1_1 GATOR2 complex protein MIOS (chains A, B)
TKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDTPYM
KCVAWYLNYDPECLLAVGEANGRVVLTSLGQDHNSKFKDLIGKEFVPKHARQCNTLAWNP
LDSDWLAAGLDKHRADFSVLIWDICVTKPLYELGQNDACLSLCWLPRDQKLLLAGMHRNL
AIFDLRNTSQKMFVNTKAVQGVTVDPYFHDRVASFYEGQVAIWDLRKFEKPVLTLTEQPK
PLTKVAWCPTRTGLLATLTRDSNIIRLYDMQHTPTPIGDETEPTIIERSVQPCDNYIASF
AWHPTSQNRMIVVTPNRTMSDFTVFERISLAWSPITSLMWACGRHLYECTEMFVARSIAA
DHKDLIHDVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHSGSVWRVTWAHPE
FGQVLASCSFDRTAAVWEEIVGESNDKLRGQSHWVKRTTLVDSRTSVTDVKFAPKHMGLM
LATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSPMIAVGSDDS
SPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAIATKDVRIFT
LKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDGCVRLWKANY
MDNWKCTGILKGNGSPVEKDIATKMRLRALSRYGLDTEQVWRNHILAGNEDPQLKSLWYT
LHFMKQYTEDMDQKSPGNKGSLVYAGIKSIVKSSLGMVESSRHNWSGLDKQSDIQNLNEE
RILALQLCGWIKKGTDVDVGPFLNSLVQEGEWERAAAVALFNLDIRRAIQILNEGASSEK
GDLNLNVVAMALSGYTDEKNSLWREMCSTLRLQLNNPYLCVMFAFLTSETGSYDGVLYEN
KVAVRDRVAFACKFLSDTQLNRYIEKLTNEMKEAGNLEGILLTGLTKDGVDLMESYVDRT
GDVQTASYCMLQGSPLDVLKDERVQYWIENYRNLLDAWRFWHKRAEFDIHRSKLDPSSKP
LAQVFVSCNFCGKSISYSKVTSCPGCRKPLPRCALCLINMGTPVSKDKKLAQFNNWFTWC
HNCRHGGHAGHMLSWFRDHAECPVSACTCKCMQLDT

Sequence of entity 2 (C), FASTA

>9DX1_2 Nucleoporin SEH1 (chains C)
ERANPEGLCYGLFGDLAFAAKESLVAEPFAGLASSALSVFETEPMFVARSIAADHKDLIH
DVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHSGSVWRVTWAHPEFGQVLAS
CSFDRTAAVWEEIVKLRGQSHWVKRTTLVDSRTSVTDVKFAPKHMGLMLATCSADGIVRI
YEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSPMIAVGSDDSSPNAMAKVQIFE
YNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAIATKDVRIFTLKPVRKELTSSG
GPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDGCVRLWKANYMDNWKCTGILKG
NGGMRWFVDTAERYALAGRPLAELCDHNAKVARELGRNQVAQTWTMLRIIYCSPDSSSFS
LLSVSHALYDSRLPPDFFGVLVRDMLHFYAEQGDVQMAVSVLIVLGERVRKDIDEQTQEH
WYTSYIDLLQRFRLWNVSNEVVKLSTSRAVSCLNQASTTLHVNCSHCKRPMSSRGWVCDR
CHRCASMCAVCHHVVKGLFVWCQGCSHGGHLQHIMKWLEGSSHCPAGCGHLCEYS

Sequence of entity 3 (D), FASTA

>9DX1_3 Protein SEC13 homolog (chains D)
ARVTTAYGSYQDANIPFPRTSGARFCGAGYLVYFTRPMTMHRAVSPTEPTPRSLSALSAY
HTRWKSKREGSDSGNRQIKAAGKVIIQDGSGSGMVSVINTVDTSHEDMIHDAQMDYYGTR
LATCSSDRSVKIFDVRNGGQILIADLRGHEGPVWQVAWAHPMYGNILASCSYDRKVIIWR
EENGTWEKSHEHAGHDSSVNSVCWAPHDYGLILACGSSDGAISLLTYTGEGQWEVKKINN
AHTIGCNAVSWAPAVVPGSLIDHPSGQKPNYIKRFASGGCDNLIKLWKEEEDGQWKEEQK
LEAHSDWVRDVAWAPSIGLPTSTIASCSQDGRVFIWTCDDASSNTWSPKLLHKFNDVVWH
VSWSITANILAVSGGDNKVTLWKESVDGQWVCISDACLLPVHKSLGELYILNVNDIQETC
QKNAASALLVGRKDLVQVWSLATVATDLCLGPKSDPDLETPWARHPFGRQLLESLLAHYC
RLRDVQTLAMLCSVFEAQSRPQGLPNLTYSDPRERERDQHDKNKRLLDPANTQQFDDFKK
CYGEILYRWGLREKRAEVLKFVSC

Sequence of entity 4 (E), FASTA

>9DX1_4 Sestrin2 (chains E)
RARRGPRGPSAFIPVEEVLRDNLAVVMGLHPDYFTSFWRLHYLLLHTDGPLASSWRHYIA
IMAAARHQCSYLVGSHMAEFLQTGGDPEWLLGLHRAPEKLRKLSEINKLLAHRPWLITKE
HIQALLKTGEHTWSLAELIQALVLLTHCHSLSSFVFGCGILPEHPDMLCFVEDPTFGYED
FTRRGAQAPPTFRAQDYTWEDHGYSLIQRLYPEGGQLLDEKFQAAYSLTYNTIAMHSGVD
TSVLRRAIWNYIHCVFGIRYDDYDYGEVNQLLERNLKVYIKTVACYPEKTTRRMYNLFWR
HFRHSEKVHVNLLLLEARMQAALLYALRAITRYMT

Sequence of entity 5 (F), FASTA

>9DX1_5 GATOR2 complex protein WDR24 (chains F)
TKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDTPYM
KCVAWYLNYDPECLLAVGEANGRVVLTSLGQDHNSKFKDLIGKEFVPKHARQCNTLAWNP
LDSDWLAAGLDKHRADFSVLIWDIPLYELGQNDACLSLCWLPRDQKLLLAGMHRNLAIFD
LRNTSQKMFVNTKAVQGVTVDPYFHDRVASFYEGQVAIWDLRKFEKPVLTLTEQPKPLTK
VAWCPTRTGLLATLTRDSNIIRLYDMQHTPTPIGDETEPTIIERSVQPCDNYIASFAWHP
TSQNRMIVVTPNRTMSDFTVFSVLTGRTMHCHLDAPANAISVCRDAAQVVVAGRSIFKIY
AIEEEQFVEKLNLRVGRKPSLNLSCADVVWHQMDENLLATAATNGVVVTWNLGRPSRNKQ
DQLFTEHKRTVNKVCFHPTEAHVLLSGSQDGFMKCFDLRRKDSVSTFSGQSESVRDVQFS
IRDYFTFASTFENGNVQLWDIRRPDRCERMFTAHNGPVFCCDWHPEDRGWLATGGRDKMV
KVWDMTTHRAKEMHCVQTIASVARVKWRPECRHHLATCSMMVDHNIYVWDVRRPFVPAAM
FEEHRDVTTGIAWRHPHDPSFLLSGSKDSSLCQHLFRDASQPVPPDPHKGIEFGVYCSHC
RSEVRGTQCAICKGFTFQCAICHVAVRGSSNFCLTCGHGGHTSHMMEWFRTQEVCPTGCG
CHCLLESTF

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:

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
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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