9LVK: CASTOR1 bound human GATOR2 complex

Cryo-EM structure of CASTOR1 bound human GATOR2 complex. Determined by electron microscopy at 3.59 Å resolution. Released 13 Aug 2025.

Method: Electron microscopy
Resolution: 3.59 Å
Organism: Homo sapiens
Chains: 18
Atoms: 51,417
Mol. weight: 1227.29 kDa
Ligands: ZN
Released: 13 Aug 2025

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

Secondary structure: helices and β-sheets

9LVK contains 223 α-helices and 535 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: 31 helices, 47 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	1
β-strand	17-21	5	1
β-strand	27-29	3	1
β-strand	44	1	2
β-strand	50	1	2
β-strand	52-55	4	1
β-strand	63-66	4	3
β-strand	77-78	2	4
β-strand	80-82	3	3
β-strand	87-88	2	5
β-strand	90-91	2	4
β-strand	107-108	2	5
β-strand	119-121	3	6
β-strand	128-131	4	6
β-strand	142	1	7
β-strand	143-146	4	6
β-strand	181	1	7
β-strand	189-192	4	8
β-strand	199-203	5	8
β-strand	208-212	5	8
β-strand	222	1	8
β-strand	228	1	9
β-strand	239-244	6	9
β-strand	247-252	6	9
β-strand	256	1	10
β-strand	258	1	10
β-strand	261-262	2	9
α-helix	263-266	4
α-helix	269	1
β-strand	270-275	6	11
β-strand	283-287	5	11
β-strand	293-296	4	11
β-strand	315-318	4	11
β-strand	327-330	4	12
β-strand	338-341	4	12
β-strand	347-350	4	12
α-helix	353-355	3
β-strand	356-359	4	13
β-strand	367-370	4	13
β-strand	373-375	3	13
β-strand	378	1	14
α-helix	390-399	10
α-helix	412-415	4
α-helix	421-438	18
α-helix	452-454	3
α-helix	456-459	4
α-helix	488-497	10
α-helix	509-515	7
α-helix	516-518	3
α-helix	522-530	9
α-helix	535-541	7
α-helix	554-561	8
α-helix	573-578	6
α-helix	580-582	3
α-helix	586-596	11
α-helix	598	1
α-helix	603-606	4
α-helix	613-622	10
α-helix	628-643	16
α-helix	646-648	3
α-helix	649-652	4
α-helix	658-668	11
α-helix	673-679	7
α-helix	693-706	14
α-helix	710-723	14
β-strand	735-736	2	15
β-strand	743	1	15
β-strand	774	1	16
β-strand	781	1	16
α-helix	782-783	2
β-strand	784	1	17
α-helix	785	1
β-strand	791	1	17
α-helix	820-822	3
β-strand	825-827	3	18
β-strand	832	1	19
β-strand	833-834	2	18
α-helix	837-845	9

Chain B: 32 helices, 42 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	155
β-strand	17-21	5	155
β-strand	25-31	7	155
β-strand	44	1	155
β-strand	50-57	8	155
β-strand	63-68	6	156
β-strand	77-82	6	156
β-strand	87-91	5	156
β-strand	107-108	2	156
β-strand	116-121	6	157
β-strand	129-133	5	157
β-strand	142-144	3	157
β-strand	179-181	3	157
β-strand	187-192	6	158
β-strand	199-204	6	158
β-strand	207-212	6	158
β-strand	219-223	5	158
β-strand	231-233	3	159
β-strand	236-241	6	159
β-strand	244	1	160
β-strand	247	1	160
β-strand	250-251	2	159
α-helix	259	1
β-strand	260	1	159
α-helix	262-265	4
β-strand	270	1	161
β-strand	285	1	162
β-strand	287	1	161
β-strand	293	1	163
β-strand	294	1	162
β-strand	296-297	2	164
α-helix	301-303	3
α-helix	309-312	4
β-strand	314-315	2	164
β-strand	318	1	163
α-helix	324-325	2
β-strand	327-330	4	165
β-strand	338-341	4	165
β-strand	347-348	2	165
β-strand	359	1	110
β-strand	366-370	5	110
β-strand	373-377	5	110
α-helix	390-399	10
α-helix	409-411	3
α-helix	412-415	4
α-helix	423-442	20
α-helix	455-460	6
α-helix	471-473	3
α-helix	488-497	10
α-helix	509-518	10
α-helix	521-530	10
α-helix	534-545	12
α-helix	554-562	9
α-helix	571-579	9
α-helix	586-595	10
β-strand	598	1	166
β-strand	601	1	166
α-helix	603-606	4
α-helix	613-623	11
α-helix	627-642	16
α-helix	646-648	3
α-helix	649-652	4
α-helix	657-669	13
α-helix	673-679	7
α-helix	691-707	17
α-helix	710-723	14
α-helix	725-727	3
β-strand	733	1	100
α-helix	791	1
α-helix	819-822	4
β-strand	824-827	4	103
β-strand	833-835	3	103
α-helix	836-843	8
α-helix	858-862	5

Chain C: 15 helices, 44 β-strands

Element	Residues	Length	Sheet
β-strand	21-24	4	20
β-strand	31-34	4	21
β-strand	40-44	5	21
β-strand	49-53	5	21
β-strand	60-61	2	21
β-strand	77-82	6	22
β-strand	89-94	6	22
β-strand	99-103	5	22
β-strand	112-116	5	22
β-strand	123-125	3	23
β-strand	128	1	23
β-strand	135-140	6	23
β-strand	145-149	5	23
β-strand	157-159	3	23
β-strand	166-171	6	24
β-strand	178-183	6	24
β-strand	187-192	6	24
β-strand	201-204	4	24
β-strand	210-215	6	25
β-strand	223-227	5	25
β-strand	231-236	6	25
β-strand	243-249	7	25
β-strand	257-259	3	26
β-strand	266-270	5	26
β-strand	278-282	5	26
β-strand	285-293	9	26
β-strand	300-305	6	20
β-strand	313-318	6	20
β-strand	322-327	6	20
α-helix	328-330	3
β-strand	332	1	26
α-helix	335-337	3
β-strand	342-344	3	27
β-strand	352-354	3	27
β-strand	356	1	28
β-strand	395	1	28
β-strand	397	1	29
α-helix	410-418	9
α-helix	426-440	15
α-helix	443-456	14
α-helix	630-632	3
α-helix	634-647	14
α-helix	650-660	11
α-helix	661-663	3
α-helix	670-686	17
α-helix	690-698	9
α-helix	703-706	4
β-strand	714	1	19
β-strand	716-717	2	15
β-strand	718	1	30
α-helix	724	1
β-strand	725	1	30
α-helix	726	1
β-strand	732	1	31
β-strand	739	1	31
β-strand	742	1	32
β-strand	749	1	32
β-strand	753-755	3	18
α-helix	766-773	8

Chain D: 13 helices, 5 β-strands

Element	Residues	Length	Sheet
β-strand	541-545	5	33
β-strand	549-553	5	33
β-strand	647-650	4	33
α-helix	660-665	6
α-helix	673-687	15
α-helix	691-701	11
α-helix	705-707	3
α-helix	726-739	14
α-helix	742-752	11
α-helix	841-848	8
α-helix	853-855	3
α-helix	856-873	18
α-helix	877-883	7
α-helix	929-931	3
β-strand	934-937	4	34
β-strand	944-946	3	34
α-helix	947-954	8
α-helix	968-970	3

Chain E: 3 helices, 29 β-strands

Element	Residues	Length	Sheet
β-strand	3	1	14
β-strand	6	1	13
β-strand	15-19	5	35
β-strand	26-31	6	35
β-strand	35-41	7	35
β-strand	47-54	8	35
β-strand	60-65	6	36
α-helix	68-70	3
β-strand	74-78	5	36
β-strand	83-86	4	36
β-strand	105	1	36
β-strand	121	1	37
α-helix	122-123	2
β-strand	125	1	38
β-strand	127	1	38
β-strand	130	1	37
β-strand	133	1	39
β-strand	138-143	6	39
β-strand	154-160	7	39
β-strand	165-170	6	40
β-strand	181-186	6	40
β-strand	197-201	5	40
β-strand	210-214	5	40
β-strand	224-227	4	41
β-strand	236-241	6	41
β-strand	245-252	8	41
α-helix	263-265	3
β-strand	266-274	9	41
β-strand	284-286	3	42
β-strand	292-295	4	42
β-strand	302-306	5	42
β-strand	314-319	6	42

Chain F: 3 helices, 36 β-strands

Element	Residues	Length	Sheet
β-strand	3-7	5	43
β-strand	26	1	44
β-strand	29	1	45
β-strand	31	1	46
β-strand	35	1	47
β-strand	37	1	45
β-strand	39-41	3	44
α-helix	42	1
β-strand	47-49	3	44
β-strand	54	1	47
β-strand	60	1	46
β-strand	64-65	2	48
α-helix	68-70	3
β-strand	73	1	49
β-strand	74-75	2	48
β-strand	77	1	50
β-strand	83	1	50
β-strand	87	1	49
β-strand	89	1	51
β-strand	102	1	51
β-strand	116-120	5	52
β-strand	129-134	6	52
β-strand	139-143	5	52
β-strand	147	1	53
β-strand	149	1	53
β-strand	154-158	5	52
β-strand	165-169	5	54
β-strand	183-186	4	54
β-strand	198-202	5	54
β-strand	209-214	6	54
α-helix	227-229	3
β-strand	236	1	55
β-strand	250	1	55
β-strand	252	1	56
β-strand	266	1	56
β-strand	284-286	3	57
β-strand	293-295	3	57
β-strand	302-303	2	58
β-strand	318-319	2	58

Chain G: 2 helices, 29 β-strands

Element	Residues	Length	Sheet
β-strand	7	1	29
β-strand	15-20	6	59
β-strand	27-31	5	59
β-strand	35-41	7	59
β-strand	47-53	7	59
β-strand	60-64	5	60
β-strand	73-78	6	60
β-strand	83-88	6	60
β-strand	103-109	7	60
β-strand	116-121	6	61
α-helix	124-126	3
β-strand	129-134	6	61
β-strand	138-143	6	61
β-strand	154-160	7	61
β-strand	165-169	5	62
α-helix	179-180	2
β-strand	181-186	6	62
β-strand	197-203	7	62
β-strand	208-214	7	62
β-strand	224-227	4	63
β-strand	236-241	6	63
β-strand	245-250	6	63
β-strand	252	1	64
β-strand	266	1	64
β-strand	269-274	6	63
β-strand	283-286	4	65
β-strand	293-296	4	65
β-strand	304-305	2	65
β-strand	307	1	66
β-strand	313	1	66
β-strand	317	1	65

Chain H: 5 helices, 28 β-strands

Element	Residues	Length	Sheet
β-strand	51-54	4	33
β-strand	62-67	6	67
β-strand	74-78	5	67
β-strand	86-87	2	67
β-strand	94-95	2	67
β-strand	106-111	6	68
α-helix	112-113	2
α-helix	114-116	3
β-strand	120-124	5	68
β-strand	129-131	3	68
β-strand	134-135	2	69
β-strand	138-139	2	69
β-strand	144-145	2	68
β-strand	152-157	6	70
α-helix	158-159	2
β-strand	165-170	6	70
β-strand	175-182	8	70
β-strand	185-188	4	70
β-strand	191	1	70
β-strand	199-204	6	71
α-helix	205-207	3
β-strand	228-232	5	71
β-strand	238-240	3	71
β-strand	250-253	4	71
β-strand	261-266	6	72
α-helix	267-269	3
β-strand	275	1	73
β-strand	277-281	5	72
β-strand	286-288	3	72
β-strand	291	1	73
β-strand	311-315	5	74
β-strand	324-327	4	74
β-strand	332-334	3	74

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, K, L	protein	875	Homo sapiens	Q9NXC5 (AlphaFold model)
GATOR2 complex protein WDR24	C, M	protein	790	Homo sapiens	Q96S15 (AlphaFold model)
GATOR2 complex protein WDR59	D, N	protein	974	Homo sapiens	Q6PJI9 (AlphaFold model)
Isoform B of Nucleoporin SEH1	E, F, G, O, P, Q	protein	421	Homo sapiens	Q96EE3 (AlphaFold model)
Isoform 3 of Protein SEC13 homolog	H, R	protein	368	Homo sapiens	P55735
Cytosolic arginine sensor for mTORC1 subunit 1	U, V	protein	329	Homo sapiens	Q8WTX7

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

>9LVK_1 GATOR2 complex protein MIOS (chains A, B, K, L)
MSGTKPDILWAPHHVDRFVVCDSELSLYHVESTVNSELKAGSLRLSEDSAATLLSINSDT
PYMKCVAWYLNYDPECLLAVGQANGRVVLTSLGQDHNSKFKDLIGKEFVPKHARQCNTLA
WNPLDSNWLAAGLDKHRADFSVLIWDICSKYTPDIVPMEKVKLSAGETETTLLVTKPLYE
LGQNDACLSLCWLPRDQKLLLAGMHRNLAIFDLRNTSQKMFVNTKAVQGVTVDPYFHDRV
ASFYEGQVAIWDLRKFEKPVLTLTEQPKPLTKVAWCPTRTGLLATLTRDSNIIRLYDMQH
TPTPIGDETEPTIIERSVQPCDNYIASFAWHPTSQNRMIVVTPNRTMSDFTVFERISLAW
SPITSLMWACGRHLYECTEEENDNSLEKDIATKMRLRALSRYGLDTEQVWRNHILAGNED
PQLKSLWYTLHFMKQYTEDMDQKSPGNKGSLVYAGIKSIVKSSLGMVESSRHNWSGLDKQ
SDIQNLNEERILALQLCGWIKKGTDVDVGPFLNSLVQEGEWERAAAVALFNLDIRRAIQI
LNEGASSEKGDLNLNVVAMALSGYTDEKNSLWREMCSTLRLQLNNPYLCVMFAFLTSETG
SYDGVLYENKVAVRDRVAFACKFLSDTQLNRYIEKLTNEMKEAGNLEGILLTGLTKDGVD
LMESYVDRTGDVQTASYCMLQGSPLDVLKDERVQYWIENYRNLLDAWRFWHKRAEFDIHR
SKLDPSSKPLAQVFVSCNFCGKSISYSCSAVPHQGRGFSQYGVSGSPTKSKVTSCPGCRK
PLPRCALCLINMGTPVSSCPGGTKSDEKVDLSKDKKLAQFNNWFTWCHNCRHGGHAGHML
SWFRDHAECPVSACTCKCMQLDTTGNLVPAETVQP

Sequence of entity 2 (C, M), FASTA

>9LVK_2 GATOR2 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

>9LVK_3 GATOR2 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

>9LVK_4 Isoform B of Nucleoporin SEH1 (chains E, F, G, O, P, Q)
MFVARSIAADHKDLIHDVSFDFHGRRMATCSSDQSVKVWDKSESGDWHCTASWKTHSGSV
WRVTWAHPEFGQVLASCSFDRTAAVWEEIVGESNDKLRGQSHWVKRTTLVDSRTSVTDVK
FAPKHMGLMLATCSADGIVRIYEAPDVMNLSQWSLQHEISCKLSCSCISWNPSSSRAHSP
MIAVGSDDSSPNAMAKVQIFEYNENTRKYAKAETLMTVTDPVHDIAFAPNLGRSFHILAI
ATKDVRIFTLKPVRKELTSSGGPTKFEIHIVAQFDNHNSQVWRVSWNITGTVLASSGDDG
CVRLWKANYMDNWKCTGILKGNGSPVNGSSQQGTSNPSLGSTIPSLQNSLNGSSAGRYFF
TPLDSPRAGSRWSSYAQLLPPPPPPLVEHSCDADTANLQYPHPRRRYLSRPLNPLPENEG
I

Sequence of entity 5 (H, R), FASTA

>9LVK_5 Isoform 3 of Protein SEC13 homolog (chains H, R)
MREPVLTWCVPLELLCSHPLPLSAFLKSQVKLYTYRACAGKDEMGKMVSVINTVDTSHED
MIHDAQMDYYGTRLATCSSDRSVKIFDVRNGGQILIADLRGHEGPVWQVAWAHPMYGNIL
ASCSYDRKVIIWREENGTWEKSHEHAGHDSSVNSVCWAPHDYGLILACGSSDGAISLLTY
TGEGQWEVKKINNAHTIGCNAVSWAPAVVPGSLIDHPSGQKPNYIKRFASGGCDNLIKLW
KEEEDGQWKEEQKLEAHSDWVRDVAWAPSIGLPTSTIASCSQDGRVFIWTCDDASSNTWS
PKLLHKFNDVVWHVSWSITANILAVSGGDNKVTLWKESVDGQWVCISDVNKGQGSVSASV
TEGQQNEQ

Sequence of entity 6 (U, V), FASTA

>9LVK_6 Cytosolic arginine sensor for mTORC1 subunit 1 (chains U, V)
MELHILEHRVRVLSVARPGLWLYTHPLIKLLFLPRRSRCKFFSLTETPEDYTLMVDEEGF
KELPPSEFLQVAEATWLVLNVSSHSGAAVQAAGVTKIARSVIAPLAEHHVSVLMLSTYQT
DFILVREQDLSVVIHTLAQEFDIYREVGGEPVPVTRDDSSNGFPRTQHGPSPTVHPIQSP
QNRFCVLTLDPETLPAIATTLIDVLFYSHSTPKEAASSSPEPSSITFFAFSLIEGYISIV
MDAETQKKFPSDLLLTSSSGELWRMVRIGGQPLGFDECGIVAQIAGPLAAADISAYYIST
FNFAHALVPEDGIGSVIEVLQRRQEGLAS

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	31

Primary citation

Cryo-EM structures of amino acid sensors bound to the human GATOR2 complex. Su, M.Y., Teng, F., Wang, S. et al. Cell Rep (2025) 44:116088-116088. DOI 10.1016/j.celrep.2025.116088 · 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
9OTI 3.5 Å, GATOR2 complex bound to arginine sensor CASTOR1
7UHY 3.66 Å, Human GATOR2 complex
9LVJ 3.82 Å, Cryo-EM structure of Sestrin2 bound human GATOR2 complex

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