9LWF: Dual sensor bound GATOR2 complex

Cryo-EM structure of dual sensor bound GATOR2 complex. Determined by electron microscopy at 3.41 Å resolution. Released 13 Aug 2025.

Method: Electron microscopy
Resolution: 3.41 Å
Organism: Homo sapiens
Chains: 20
Atoms: 57,308
Mol. weight: 1336.45 kDa
Ligands: ZN
Released: 13 Aug 2025

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

Secondary structure: helices and β-sheets

9LWF contains 266 α-helices and 552 β-strands across 20 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: 30 helices, 42 β-strands

Element	Residues	Length	Sheet
β-strand	8-10	3	1
β-strand	17-20	4	1
β-strand	25-29	5	1
β-strand	52-57	6	1
β-strand	63-68	6	2
β-strand	77-82	6	2
β-strand	87	1	2
β-strand	90-91	2	2
β-strand	118-121	4	3
β-strand	128-132	5	3
β-strand	142-146	5	3
β-strand	179-181	3	3
β-strand	189-192	4	4
β-strand	199-203	5	4
β-strand	207-212	6	4
β-strand	222-223	2	4
β-strand	228	1	5
β-strand	239-244	6	5
β-strand	247-252	6	5
β-strand	256	1	6
β-strand	258	1	6
β-strand	261-262	2	5
α-helix	263-266	4
α-helix	269	1
β-strand	270-275	6	7
β-strand	282-287	6	7
β-strand	293-297	5	7
β-strand	315-318	4	7
β-strand	327-330	4	8
β-strand	337-341	5	8
β-strand	347-351	5	8
α-helix	353-355	3
β-strand	356-360	5	9
β-strand	366-370	5	9
β-strand	373-378	6	9
α-helix	390-398	9
α-helix	409-411	3
α-helix	412-415	4
α-helix	421-438	18
α-helix	456-461	6
α-helix	489-497	9
α-helix	509-517	9
α-helix	522-530	9
α-helix	534-538	5
α-helix	554-557	4
α-helix	560-562	3
α-helix	571-576	6
α-helix	580-582	3
α-helix	586-595	10
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	693-706	14
α-helix	710-723	14
β-strand	733-735	3	10
β-strand	736	1	11
β-strand	743	1	11
β-strand	774	1	12
β-strand	781	1	12
α-helix	782-783	2
β-strand	784	1	13
α-helix	785	1
β-strand	791	1	13
α-helix	820-822	3
β-strand	824-827	4	14
β-strand	832	1	15
β-strand	833-835	3	14
α-helix	836-845	10

Chain B: 25 helices, 43 β-strands

Element	Residues	Length	Sheet
β-strand	7-10	4	16
β-strand	18-21	4	16
β-strand	25-27	3	16
β-strand	30-31	2	17
β-strand	43-44	2	17
β-strand	50-51	2	17
β-strand	55-57	3	16
β-strand	63-65	3	18
β-strand	68	1	18
β-strand	77-82	6	18
β-strand	87-91	5	18
β-strand	108	1	18
β-strand	112	1	19
β-strand	116-119	4	20
β-strand	130-133	4	20
β-strand	140-144	5	20
β-strand	174	1	18
β-strand	179-183	5	20
β-strand	187-192	6	21
β-strand	199-204	6	21
β-strand	208-212	5	21
β-strand	221-222	2	21
β-strand	231-232	2	22
β-strand	240-244	5	22
β-strand	247-250	4	22
β-strand	260	1	22
β-strand	270-275	6	23
β-strand	283-287	5	23
β-strand	293	1	24
β-strand	296	1	25
β-strand	315	1	25
β-strand	318	1	24
β-strand	328-329	2	26
β-strand	339-340	2	26
β-strand	356-359	4	27
β-strand	366-370	5	27
β-strand	373-377	5	27
α-helix	390-398	9
α-helix	409-411	3
α-helix	412-415	4
α-helix	423-429	7
α-helix	456-459	4
α-helix	493-496	4
α-helix	509-515	7
α-helix	516-518	3
α-helix	522-529	8
α-helix	534-542	9
α-helix	556-561	6
α-helix	571-579	9
α-helix	586-593	8
α-helix	613-623	11
α-helix	626-639	14
α-helix	646-648	3
α-helix	649-652	4
α-helix	658-669	12
α-helix	673-677	5
α-helix	693-706	14
α-helix	710-723	14
α-helix	729-731	3
β-strand	733	1	28
β-strand	784	1	29
α-helix	790	1
β-strand	791	1	29
β-strand	824	1	30
β-strand	826-827	2	31
β-strand	835	1	30
α-helix	836-845	10
α-helix	859-861	3

Chain C: 11 helices, 41 β-strands

Element	Residues	Length	Sheet
β-strand	22-24	3	32
β-strand	31-34	4	33
β-strand	40-44	5	33
β-strand	49-54	6	33
β-strand	59-64	6	33
β-strand	79-82	4	34
β-strand	89-93	5	34
β-strand	99-103	5	34
β-strand	112-116	5	34
β-strand	123-128	6	35
β-strand	135-140	6	35
β-strand	144-149	6	35
β-strand	159-160	2	35
β-strand	166-168	3	36
β-strand	171	1	36
β-strand	179-183	5	36
β-strand	187-191	5	36
β-strand	201-204	4	36
β-strand	210-215	6	37
β-strand	222-227	6	37
β-strand	231-236	6	37
β-strand	243-249	7	37
β-strand	254-259	6	38
α-helix	260	1
β-strand	267-271	5	38
β-strand	278-280	3	38
β-strand	291-293	3	38
β-strand	300-305	6	32
β-strand	314-318	5	32
β-strand	322-326	5	32
α-helix	328-330	3
β-strand	332	1	38
β-strand	342-345	4	39
β-strand	351-356	6	39
β-strand	395-399	5	39
β-strand	400	1	40
α-helix	411-416	6
α-helix	426-440	15
α-helix	443-456	14
α-helix	634-646	13
α-helix	650-660	11
α-helix	672-684	13
α-helix	690-698	9
α-helix	708-710	3
β-strand	714	1	41
β-strand	717-719	3	42
β-strand	725	1	42
β-strand	732	1	43
β-strand	739	1	43
β-strand	753-756	4	44
β-strand	763-765	3	44
α-helix	769-772	4

Chain D: 14 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	536-538	3
β-strand	541-545	5	45
β-strand	549-553	5	45
β-strand	647-650	4	45
α-helix	660-665	6
α-helix	673-687	15
α-helix	690-701	12
α-helix	705-707	3
α-helix	726-739	14
α-helix	742-751	10
α-helix	839-847	9
α-helix	856-873	18
α-helix	877-883	7
β-strand	900-901	2	46
α-helix	902	1
β-strand	908-909	2	46
β-strand	913	1	47
β-strand	920	1	47
β-strand	923	1	48
β-strand	930	1	48
α-helix	931	1
β-strand	934-937	4	49
β-strand	944-946	3	49
α-helix	947-953	7
α-helix	968-971	4

Chain E: 3 helices, 27 β-strands

Element	Residues	Length	Sheet
β-strand	3-7	5	9
β-strand	15-20	6	50
β-strand	26-31	6	50
β-strand	35-40	6	50
β-strand	48-54	7	50
β-strand	60-64	5	51
α-helix	68-70	3
β-strand	73-78	6	51
β-strand	83-89	7	51
β-strand	102-109	8	51
β-strand	116-121	6	52
α-helix	122-123	2
α-helix	124-126	3
β-strand	129-134	6	52
β-strand	138-143	6	52
β-strand	154-160	7	52
β-strand	165-170	6	53
β-strand	181-186	6	53
β-strand	197-203	7	53
β-strand	208-214	7	53
β-strand	224-227	4	54
β-strand	236-241	6	54
β-strand	245-250	6	54
β-strand	252	1	55
β-strand	266	1	55
β-strand	269-274	6	54
β-strand	283-286	4	56
β-strand	293-296	4	56
β-strand	301-306	6	56
β-strand	314-320	7	56

Chain F: 4 helices, 34 β-strands

Element	Residues	Length	Sheet
β-strand	3	1	57
β-strand	18-20	3	58
β-strand	27-31	5	58
β-strand	35-41	7	58
β-strand	47-53	7	58
β-strand	60-65	6	59
α-helix	68-70	3
β-strand	74-78	5	59
β-strand	82-86	5	59
β-strand	89	1	60
β-strand	102	1	60
β-strand	105	1	59
β-strand	116-118	3	61
β-strand	121	1	62
α-helix	124-126	3
β-strand	130	1	62
β-strand	133-134	2	61
β-strand	138-139	2	61
β-strand	142	1	62
β-strand	159-160	2	61
β-strand	167-169	3	63
β-strand	182-185	4	63
β-strand	197-200	4	63
β-strand	211-214	4	63
α-helix	221-222	2
β-strand	225	1	64
α-helix	227-229	3
β-strand	236	1	65
β-strand	239-241	3	64
β-strand	245-247	3	64
β-strand	250	1	65
β-strand	251-252	2	66
β-strand	266-267	2	66
β-strand	272-274	3	64
β-strand	284-286	3	67
β-strand	293-295	3	67
β-strand	303-306	4	67
β-strand	314-315	2	67

Chain G: 5 helices, 25 β-strands

Element	Residues	Length	Sheet
β-strand	4-7	4	68
β-strand	15-20	6	69
β-strand	26-31	6	69
β-strand	35-41	7	69
β-strand	47-54	8	69
β-strand	60-65	6	70
β-strand	73-78	6	70
β-strand	83-89	7	70
β-strand	102-109	8	70
β-strand	116-121	6	71
α-helix	122-123	2
α-helix	124-126	3
β-strand	129-134	6	71
β-strand	138-143	6	71
β-strand	154-160	7	71
β-strand	165-170	6	72
β-strand	181-186	6	72
β-strand	197-203	7	72
β-strand	208-214	7	72
α-helix	221-222	2
β-strand	224-227	4	73
β-strand	236-241	6	73
β-strand	245-252	8	73
α-helix	253-254	2
α-helix	263-265	3
β-strand	266-274	9	73
β-strand	281-286	6	74
β-strand	293-297	5	74
β-strand	301-307	7	74
β-strand	313-320	8	74

Chain H: 4 helices, 27 β-strands

Element	Residues	Length	Sheet
β-strand	54	1	45
β-strand	65-67	3	75
β-strand	74-77	4	75
β-strand	82-87	6	75
β-strand	94-100	7	75
β-strand	106-111	6	76
α-helix	112-113	2
α-helix	114-116	3
β-strand	120-124	5	76
β-strand	129-135	7	76
β-strand	138-145	8	76
β-strand	152-157	6	77
α-helix	158-159	2
β-strand	165-170	6	77
β-strand	175-180	6	77
β-strand	186-191	6	77
β-strand	199-204	6	78
α-helix	205-207	3
β-strand	228-232	5	78
β-strand	237-242	6	78
β-strand	248-254	7	78
β-strand	261-266	6	79
β-strand	275	1	80
β-strand	277-281	5	79
β-strand	290	1	81
β-strand	291	1	80
β-strand	300	1	81
β-strand	311-315	5	82
β-strand	322-327	6	82
β-strand	332-337	6	82
β-strand	343-345	3	82

12 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, I	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
Sestrin-2	X, Y	protein	480	Homo sapiens	P58004

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

>9LWF_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, I), FASTA

>9LWF_2 GATOR2 complex protein WDR24 (chains C, I)
MEKMSRVTTALGGSVLTGRTMHCHLDAPANAISVCRDAAQVVVAGRSIFKIYAIEEEQFV
EKLNLRVGRKPSLNLSCADVVWHQMDENLLATAATNGVVVTWNLGRPSRNKQDQLFTEHK
RTVNKVCFHPTEAHVLLSGSQDGFMKCFDLRRKDSVSTFSGQSESVRDVQFSIRDYFTFA
STFENGNVQLWDIRRPDRCERMFTAHNGPVFCCDWHPEDRGWLATGGRDKMVKVWDMTTH
RAKEMHCVQTIASVARVKWRPECRHHLATCSMMVDHNIYVWDVRRPFVPAAMFEEHRDVT
TGIAWRHPHDPSFLLSGSKDSSLCQHLFRDASQPVERANPEGLCYGLFGDLAFAAKESLV
AAESGRKPYTGDRRHPIFFKRKLDPAEPFAGLASSALSVFETEPGGGGMRWFVDTAERYA
LAGRPLAELCDHNAKVARELGRNQVAQTWTMLRIIYCSPGLVPTANLNHSVGKGGSCGLP
LMNSFNLKDMAPGLGSETRLDRSKGDARSDTVLLDSSATLITNEDNEETEGSDVPADYLL
GDVEGEEDELYLLDPEHAHPEDPECVLPQEAFPLRHEIVDTPPGPEHLQDKADSPHVSGS
EADVASLAPVDSSFSLLSVSHALYDSRLPPDFFGVLVRDMLHFYAEQGDVQMAVSVLIVL
GERVRKDIDEQTQEHWYTSYIDLLQRFRLWNVSNEVVKLSTSRAVSCLNQASTTLHVNCS
HCKRPMSSRGWVCDRCHRCASMCAVCHHVVKGLFVWCQGCSHGGHLQHIMKWLEGSSHCP
AGCGHLCEYS

Sequence of entity 3 (D, N), FASTA

>9LWF_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

>9LWF_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

>9LWF_5 Isoform 3 of Protein SEC13 homolog (chains H, R)
MREPVLTWCVPLELLCSHPLPLSAFLKSQVKLYTYRACAGKDEMGKMVSVINTVDTSHED
MIHDAQMDYYGTRLATCSSDRSVKIFDVRNGGQILIADLRGHEGPVWQVAWAHPMYGNIL
ASCSYDRKVIIWREENGTWEKSHEHAGHDSSVNSVCWAPHDYGLILACGSSDGAISLLTY
TGEGQWEVKKINNAHTIGCNAVSWAPAVVPGSLIDHPSGQKPNYIKRFASGGCDNLIKLW
KEEEDGQWKEEQKLEAHSDWVRDVAWAPSIGLPTSTIASCSQDGRVFIWTCDDASSNTWS
PKLLHKFNDVVWHVSWSITANILAVSGGDNKVTLWKESVDGQWVCISDVNKGQGSVSASV
TEGQQNEQ

Sequence of entity 6 (U, V), FASTA

>9LWF_6 Cytosolic arginine sensor for mTORC1 subunit 1 (chains U, V)
MELHILEHRVRVLSVARPGLWLYTHPLIKLLFLPRRSRCKFFSLTETPEDYTLMVDEEGF
KELPPSEFLQVAEATWLVLNVSSHSGAAVQAAGVTKIARSVIAPLAEHHVSVLMLSTYQT
DFILVREQDLSVVIHTLAQEFDIYREVGGEPVPVTRDDSSNGFPRTQHGPSPTVHPIQSP
QNRFCVLTLDPETLPAIATTLIDVLFYSHSTPKEAASSSPEPSSITFFAFSLIEGYISIV
MDAETQKKFPSDLLLTSSSGELWRMVRIGGQPLGFDECGIVAQIAGPLAAADISAYYIST
FNFAHALVPEDGIGSVIEVLQRRQEGLAS

Sequence of entity 7 (X, Y), FASTA

>9LWF_7 Sestrin-2 (chains X, Y)
MIVADSECRAELKDYLRFAPGGVGDSGPGEEQRESRARRGPRGPSAFIPVEEVLREGAES
LEQHLGLEALMSSGRVDNLAVVMGLHPDYFTSFWRLHYLLLHTDGPLASSWRHYIAIMAA
ARHQCSYLVGSHMAEFLQTGGDPEWLLGLHRAPEKLRKLSEINKLLAHRPWLITKEHIQA
LLKTGEHTWSLAELIQALVLLTHCHSLSSFVFGCGILPEGDADGSPAPQAPTPPSEQSSP
PSRDPLNNSGGFESARDVEALMERMQQLQESLLRDEGTSQEEMESRFELEKSESLLVTPS
ADILEPSPHPDMLCFVEDPTFGYEDFTRRGAQAPPTFRAQDYTWEDHGYSLIQRLYPEGG
QLLDEKFQAAYSLTFNTIAMHSGVDTSVLRRAIWNYIHCVFGIRYDDYDYGEVNQLLERN
LKVYIKTVACYPEKTTRRMYNLFWRHFRHSEKVHVNLLLLEARMQAALLYALRAITRYMT

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	32

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
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

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