9O5D: The KICSTOR-GATOR1-SAMTOR complex

The KICSTOR-GATOR1-SAMTOR complex. Determined by electron microscopy at 3.34 Å resolution. Released 21 Jan 2026.

Method: Electron microscopy
Resolution: 3.34 Å
Organism: Homo sapiens
Chains: 5
Atoms: 22,702
Mol. weight: 491.76 kDa
Released: 21 Jan 2026

Explore 9O5D in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9O5D contains 148 α-helices and 145 β-strands across 5 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: 10 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	95-106	12
β-strand	166-171	6	21
α-helix	177-179	3
β-strand	184-190	7	21
β-strand	199-200	2	21
β-strand	201	1	22
α-helix	203-205	3
β-strand	207-209	3	23
α-helix	211	1
α-helix	216-224	9
β-strand	230	1	22
β-strand	232-234	3	23
β-strand	238-244	7	21
α-helix	247-249	3
α-helix	253-266	14
β-strand	267-278	12	21
α-helix	287-298	12
β-strand	302-310	9	21
β-strand	313-320	8	21
α-helix	335-337	3
α-helix	341-343	3

Chain B: 49 helices, 55 β-strands

Element	Residues	Length	Sheet
β-strand	38-40	3	9
β-strand	52-54	3	9
α-helix	71-76	6
β-strand	88-89	2	9
β-strand	96	1	10
β-strand	99-104	6	11
α-helix	111-121	11
β-strand	124-127	4	10
β-strand	130-134	5	11
β-strand	137-146	10	11
β-strand	149-151	3	11
β-strand	153-156	4	10
β-strand	161-166	6	11
β-strand	170-176	7	12
α-helix	179-182	4
β-strand	184	1	13
β-strand	190	1	13
α-helix	191-193	3
α-helix	194-200	7
α-helix	201-209	9
β-strand	215-226	12	12
α-helix	231-233	3
α-helix	236-238	3
β-strand	243-244	2	12
β-strand	250-262	13	12
α-helix	269-284	16
α-helix	294-295	2
β-strand	296-298	3	12
α-helix	299-300	2
α-helix	306-317	12
β-strand	322	1	14
β-strand	331-337	7	12
β-strand	342-345	4	15
α-helix	347-360	14
β-strand	363-368	6	12
α-helix	371-372	2
β-strand	379-382	4	15
β-strand	397-398	2	15
β-strand	404-407	4	12
α-helix	424-427	4
β-strand	428	1	4
α-helix	429-430	2
α-helix	459-466	8
α-helix	613-620	8
α-helix	655-663	9
α-helix	729	1
α-helix	731-734	4
α-helix	735-737	3
α-helix	744-748	5
α-helix	750	1
β-strand	751	1	11
α-helix	752-754	3
β-strand	756-757	2	16
α-helix	760-762	3
α-helix	763-768	6
β-strand	770-777	8	16
α-helix	792-794	3
α-helix	795-809	15
β-strand	812-814	3	16
β-strand	851-856	6	16
β-strand	859-865	7	16
β-strand	870-877	8	16
β-strand	886-893	8	12
β-strand	902-908	7	12
α-helix	918-926	9
α-helix	931-933	3
α-helix	934-936	3
α-helix	937-939	3
β-strand	942-950	9	14
α-helix	953-960	8
α-helix	979-994	16
α-helix	1177-1184	8
α-helix	1193	1
β-strand	1194	1	17
α-helix	1195-1196	2
β-strand	1197	1	18
β-strand	1200	1	18
β-strand	1204-1206	3	17
α-helix	1207-1215	9
α-helix	1225-1236	12
β-strand	1241-1242	2	17
β-strand	1254-1255	2	19
β-strand	1257-1259	3	17
β-strand	1296-1298	3	19
α-helix	1310-1312	3
β-strand	1351-1357	7	14
β-strand	1369-1375	7	14
β-strand	1377-1378	2	19
β-strand	1382-1383	2	20
β-strand	1384-1390	7	14
α-helix	1395-1411	17
β-strand	1414-1419	6	14
β-strand	1439-1442	4	20
α-helix	1445-1447	3
α-helix	1462-1464	3
α-helix	1465-1479	15
β-strand	1482-1485	4	20
β-strand	1501-1504	4	20
β-strand	1509-1512	4	20
β-strand	1545-1550	6	20
α-helix	1556-1558	3
α-helix	1568-1580	13
α-helix	1583-1585	3
α-helix	1586-1597	12

Chain C: 23 helices, 19 β-strands

Element	Residues	Length	Sheet
β-strand	9-15	7	1
β-strand	21-26	6	1
β-strand	47	1	11
α-helix	66-73	8
α-helix	77-79	3
β-strand	84-88	5	1
β-strand	91-95	5	1
β-strand	98-99	2	1
β-strand	123-131	9	1
α-helix	136-160	25
α-helix	162-179	18
α-helix	191-194	4
α-helix	199-213	15
β-strand	216-221	6	24
β-strand	225-230	6	24
α-helix	232-235	4
β-strand	237	1	25
β-strand	242	1	25
α-helix	245-252	8
β-strand	260-263	4	26
α-helix	267-272	6
α-helix	280-288	9
β-strand	294	1	26
α-helix	295-302	8
α-helix	306-318	13
β-strand	322-325	4	26
α-helix	326-328	3
β-strand	333-336	4	27
α-helix	347-355	9
α-helix	361-367	7
β-strand	372-373	2	27
α-helix	374-377	4
α-helix	388-399	12
β-strand	403-412	10	27
α-helix	481-487	7
α-helix	492-499	8
α-helix	502-505	4
α-helix	507-516	10
β-strand	525	1	27
α-helix	526-533	8
α-helix	537-546	10
β-strand	551-557	7	27

Chain D: 24 helices, 20 β-strands

Element	Residues	Length	Sheet
β-strand	9-16	8	1
β-strand	20-27	8	1
α-helix	34-44	11
α-helix	48-50	3
β-strand	55-59	5	1
β-strand	62-66	5	1
β-strand	69-71	3	1
β-strand	81-90	10	1
α-helix	98-116	19
α-helix	122-126	5
α-helix	128-142	15
β-strand	144-148	5	2
β-strand	154-158	5	2
α-helix	163-170	8
β-strand	174-177	4	3
α-helix	181-183	3
β-strand	184	1	4
α-helix	186-188	3
α-helix	191-196	6
α-helix	197-199	3
α-helix	206-213	8
α-helix	217-229	13
β-strand	233-236	4	3
α-helix	237-239	3
β-strand	244	1	5
β-strand	245-247	3	6
α-helix	249-251	3
α-helix	252-256	5
α-helix	258-268	11
β-strand	269	1	7
α-helix	270	1
α-helix	274-277	4
α-helix	278-287	10
β-strand	293	1	5
α-helix	294-300	7
α-helix	304-306	3
β-strand	308	1	7
α-helix	310-319	10
β-strand	323-325	3	6
β-strand	328-332	5	8
β-strand	346	1	8
α-helix	351-358	8
α-helix	362-369	8
β-strand	375-379	5	8

Chain E: 42 helices, 40 β-strands

Element	Residues	Length	Sheet
α-helix	11	1
β-strand	12	1	28
α-helix	13	1
β-strand	15-20	6	29
α-helix	28-37	10
β-strand	42-44	3	28
α-helix	45-47	3
α-helix	48-51	4
β-strand	58-64	7	29
β-strand	78-80	3	28
β-strand	86-90	5	29
β-strand	93-101	9	30
β-strand	106	1	31
β-strand	108	1	31
β-strand	109-110	2	32
β-strand	115-116	2	32
α-helix	117-133	17
β-strand	136-137	2	33
α-helix	138	1
β-strand	144-145	2	33
β-strand	148-156	9	30
β-strand	167-174	8	30
α-helix	176-178	3
α-helix	179-202	24
α-helix	235-248	14
β-strand	256-263	8	30
α-helix	272-284	13
β-strand	287-293	7	30
α-helix	310-319	10
β-strand	324-325	2	30
α-helix	340-341	2
α-helix	342-346	5
β-strand	348	1	29
β-strand	351	1	34
α-helix	374-378	5
β-strand	386-395	10	34
α-helix	399-408	10
β-strand	412-417	6	34
β-strand	424-434	11	34
β-strand	437-446	10	34
α-helix	449-450	2
β-strand	457-465	9	34
α-helix	467-474	8
α-helix	484-515	32
α-helix	517-519	3
α-helix	522-523	2
α-helix	524-528	5
β-strand	533-534	2	35
β-strand	543-544	2	35
α-helix	553-567	15
α-helix	573-576	4
β-strand	577-586	10	35
α-helix	587-589	3
α-helix	592-593	2
α-helix	611-624	14
β-strand	627-630	4	35
β-strand	634-639	6	35
α-helix	646-647	2
β-strand	649-658	10	35
β-strand	661-669	9	35
α-helix	674-688	15
β-strand	739-742	4	35
α-helix	746-749	4
α-helix	764-767	4
α-helix	782-792	11
β-strand	794-801	8	36
α-helix	812-828	17
α-helix	831	1
β-strand	832-837	6	36
β-strand	840-848	9	36
β-strand	864-871	8	36
α-helix	872-874	3
β-strand	875-876	2	36
β-strand	902-910	9	36
α-helix	911-912	2
α-helix	922-927	6
α-helix	932-934	3
α-helix	936-961	26
β-strand	986-990	5	36
α-helix	995-998	4
α-helix	1050-1052	3
α-helix	1053-1064	12

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
GATOR complex protein NPRL2	D	protein	401	Homo sapiens	Q8WTW4 (AlphaFold model)
GATOR1 complex protein DEPDC5	B	protein	1621	Homo sapiens	O75140 (AlphaFold model)
S-adenosylmethionine sensor upstream of mTORC1	A	protein	420	Homo sapiens	Q1RMZ1 (AlphaFold model)
GATOR complex protein NPRL3	C	protein	590	Homo sapiens	Q12980 (AlphaFold model)
KICSTOR complex protein SZT2	E	protein	1349	Homo sapiens	Q5T011

Sequence of entity 1 (D), FASTA

>9O5D_1 GATOR complex protein NPRL2 (chains D)
MGYPYDVPDYADLNGGGGGSTMGSGCRIECIFFSEFHPTLGPKITYQVPEDFISRELFDT
VQVYIITKPELQNKLITVTAMEKKLIGCPVCIEHKKYSRNALLFNLGFVCDAQAKTCALE
PIVKKLAGYLTTLELESSFVSMEESKQKLVPIMTILLEELNASGRCTLPIDESNTIHLKV
IEQRPDPPVAQEYDVPVFTKDKEDFFNSQWDLTTQQILPYIDGFRHIQKISAEADVELNL
VRIAIQNLLYYGVVTLVSILQYSNVYCPTPKVQDLVDDKSLQEACLSYVTKQGHKRASLR
DVFQLYCSLSPGTTVRDLIGRHPQQLQHVDERKLIQFGLMKNLIRRLQKYPVRVTREEQS
HPARLYTGCHSYDEICCKTGMSYHELDERLENDPNIIICWK

Sequence of entity 2 (B), FASTA

>9O5D_2 GATOR1 complex protein DEPDC5 (chains B)
MGDYKDDDDKGGGGGASTMRTTKVYKLVIHKKGFGGSDDELVVNPKVFPHIKLGDIVEIA
HPNDEYSPLLLQVKSLKEDLQKETISVDQTVTQVFRLRPYQDVYVNVVDPKDVTLDLVEL
TFKDQYIGRGDMWRLKKSLVSTCAYITQKVEFAGIRAQAGELWVKNEKVMCGYISEDTRV
VFRSTSAMVYIFIQMSCEMWDFDIYGDLYFEKAVNGFLADLFTKWKEKNCSHEVTVVLFS
RTFYDAKSVDEFPEINRASIRQDHKGRFYEDFYKVVVQNERREEWTSLLVTIKKLFIQYP
VLVRLEQAEGFPQGDNSTSAQGNYLEAINLSFNVFDKHYINRNFDRTGQMSVVITPGVGV
FEVDRLLMILTKQRMIDNGIGVDLVCMGEQPLHAVPLFKLHNRSAPRDSRLGDDYNIPHW
INHSFYTSKSQLFCNSFTPRIKLAGKKPASEKAKNGRDTSLGSPKESENALPIQVDYDAY
DAQVFRLPGPSRAQCLTTCRSVRERESHSRKSASSCDVSSSPSLPSRTLPTEEVRSQASD
DSSLGKSANILMIPHPHLHQYEVSSSLGYTSTRDVLENMMEPPQRDSSAPGRFHVGSAES
MLHVRPGGYTPQRALINPFAPSRMPMKLTSNRRRWMHTFPVGPSGEAIQIHHQTRQNMAE
LQGSGQRDPTHSSAELLELAYHEAAGRHSNSRQPGDGMSFLNFSGTEELSVGLLSNSGAG
MNPRTQNKDSLEDSVSTSPDPILTLSAPPVVPGFCCTVGVDWKSLTTPACLPLTTDYFPD
RQGLQNDYTEGCYDLLPEADIDRRDEDGVQMTAQQVFEEFICQRLMQGYQIIVQPKTQKP
NPAVPPPLSSSPLYSRGLVSRNRPEEEDQYWLSMGRTFHKVTLKDKMITVTRYLPKYPYE
SAQIHYTYSLCPSHSDSEFVSCWVEFSHERLEEYKWNYLDQYICSAGSEDFSLIESLKFW
RTRFLLLPACVTATKRITEGEAHCDIYGDRPRADEDEWQLLDGFVRFVEGLNRIRRRHRS
DRMMRKGTAMKGLQMTGPISTHSLESTAPPVGKKGTSALSALLEMEASQKCLGEQQAAVH
GGKSSAQSAESSSVAMTPTYMDSPRKDGAFFMEFVRSPRTASSAFYPQVSVDQTATPMLD
GTSLGICTGQSMDRGNSQTFGNSQNIGEQGYSSTNSSDSSSQQLVASSLTSSSTLTEILE
AMKHPSTGVQLLSEQKGLSPYCFISAEVVHWLVNHVEGIQTQAMAIDIMQKMLEEQLITH
ASGEAWRTFIYGFYFYKIVTDKEPDRVAMQQPATTWHTAGVDDFASFQRKWFEVAFVAEE
LVHSEIPAFLLPWLPSRPASYASRHSSFSRSFGGRSQAAALLAATVPEQRTVTLDVDVNN
RTDRLEWCSCYYHGNFSLNAAFEIKLHWMAVTAAVLFEMVQGWHRKATSCGFLLVPVLEG
PFALPSYLYGDPLRAQLFIPLNISCLLKEGSEHLFDSFEPETYWDRMHLFQEAIAHRFGF
VQDKYSASAFNFPAENKPQYIHVTGTVFLQLPYSKRKFSGQQRRRRNSTSSTNQNMFCEE
RVGYNWAYNTMLTKTWRSSATGDEKFADRLLKDFTDFCINRDNRLVTFWTSCLEKMHASA
P

Sequence of entity 3 (A), FASTA

>9O5D_3 S-adenosylmethionine sensor upstream of mTORC1 (chains A)
MEPGAGGRNTARAQRAGSPNTPPPREQERKLEQEKLSGVVKSVHRRLRKKYREVGDFDKI
WREHCEDEETLCEYAVAMKNLADNHWAKTCEGEGRIEWCCSVCREYFQNGGKRKALEKDE
KRAVLATKTTPALNMHESSQLEGHLTNLSFTNPEFITELLQASGKIRLLDVGSCFNPFLK
FEEFLTVGIDIVPAVESVYKCDFLNLQLQQPLQLAQDAIDAFLKQLKNPIDSLPGELFHV
VVFSLLLSYFPSPYQRWICCKKAHELLVLNGLLLIITPDSSHQNRHAMMMKSWKIAIESL
GFKRFKYSKFSHMHLMAFRKISLKTTSDLVSRNYPGMLYIPQDFNSIEDEEYSNPSCYVR
SDIEDEQLAYGFTELPDAPYDSDSGESQASSIPFYELEDPILLLSGGGSGGGDYKDDDDK

Sequence of entity 4 (C), FASTA

>9O5D_4 GATOR complex protein NPRL3 (chains C)
MGYPYDVPDYADLNGGGGGSTMRDNTSPISVILVSSGSRGNKLLFRYPFQRSQEHPASQT
SKPRSRYAASNTGDHADEQDGDSRFSDVILATILATKSEMCGQKFELKIDNVRFVGHPTL
LQHALGQISKTDPSPKREAPTMILFNVVFALRANADPSVINCLHNLSRRIATVLQHEERR
CQYLTREAKLILALQDEVSAMADGNEGPQSPFHHILPKCKLARDLKEAYDSLCTSGVVRL
HINSWLEVSFCLPHKIHYAASSLIPPEAIERSLKAIRPYHALLLLSDEKSLLGELPIDCS
PALVRVIKTTSAVKNLQQLAQDADLALLQVFQLAAHLVYWGKAIIIYPLCENNVYMLSPN
ASVCLYSPLAEQFSHQFPSHDLPSVLAKFSLPVSLSEFRNPLAPAVQETQLIQMVVWMLQ
RRLLIQLHTYVCLMASPSEEEPRPREDDVPFTARVGGRSLSTPNALSFGSPTSSDDMTLT
SPSMDNSSAELLPSGDSPLNQRMTENLLASLSEHERAAILSVPAAQNPEDLRMFARLLHY
FRGRHHLEEIMYNENTRRSQLLMLFDKFRSVLVVTTHEDPVIAVFQALLP

Sequence of entity 5 (E), FASTA

>9O5D_5 KICSTOR complex protein SZT2 (chains E)
MEMDYKDDDDKGGGGGASTMASERPEPEVEEAGQVFLLMKKDYRISRNVRLAWFLSHLHQ
TVQATPQEMLLQSEQELEVLSVLPPGWQPDEPVVPRPFLLVPSTRVTFLAWQYRFVIELD
LSPSTGIVDDSTGEILFDEVFHALSRCLGGLLRPFRVPGSCIDFQPEIYVTIQAYSSIIG
LQSHQVLVQGCLLDPSQREVFLQQIYEQLCLFEDKVATMLQQQYDPQSQAEDQSPDSGDL
LGRKVGVSMVTADLGLVSMIRQGILALQLLPSNSSAGIIVITDGVTSVPDVAVCETLLNQ
LRSGTVACSFVQVGGVYSYDCSFGHVPNVELMKFIAMATFGSYLSTCPEPEPGNLGLTVY
HRAFLLYSFLRSGEALNPEYYCGSQHRLFNEHLVSASSNPALALRRKKHTEKEVPADLVS
TVSVRLREGYSVREVTLAKGGSQLEVKLVLLWKHNMRIEYVAMAPWPLEPEGPRVTRVEV
TMEGGYDILHDVSCALRQPIRSLYRTHVIRRFWNTLQSINQTDQMLAHLQSFSSVPEHFT
LPDSTKSGVPLFYIPPGSTTPVLSLQPSGSDSSHAQFAAYWKPVLSMDANSWQRWLHMHR
LVLILEHDTPIPKHLHTPGSNGRYSTIQCRISHSSLTSLLRDWSSFVLVEGYSYVKLLSS
APDQPPNSFYMVRIISKAPCMVLRLGFPIGTPAPARHKIVSGLREEILRLRFPHRVQSKE
PTPKVKRKGLGGAGGGSSPSKSPPVLGPQQALSDRPCLVVLHKPLDKLLIRYEKLPLDYR
APFLLTLEPPGPLPLVSGRSASSSLASLSRYLYHQRWLWSVPSGLAPALPLSAIAQLLSI
LTEVRLSEGFHFACSGEGIINMVLELPIQNEPPGQAAAEEKHTCVVQYILFPPHSTSTKD
SFSTDDDNDVEVEALEGDSELNLVTEVWVEPQYGRVGPGPGIWKHLQDLTYSEIPQALHP
RDAACIGSMLSFEYLIQLCQSKEWGPLPPEPRVSDGLDQGGDTCVHEIPFHFDLMGLLPQ
CQQLQMFFLLLAREPEGVPFAEGSCPANDMVLCLLHSCLGQELSDREIPLTPVDQAAFLS
EVLRRTCHVPGAEGPLLGVHGIPKEQAVGSTQATGDSAFTSLSVGLPETLKPLISAQPPQ
WRCYARLVNPQHVFLTFLPATFSDVQRLAACGLEGPPQEETKPKFGDWSGAPSLKDLGGT
GIKATKSHVPVLSVTLASDNAQNQGELSPPFRRDLQAYAGRQASQTESADGPRTRCPVYI
YSCSLEALREQMVGMQPPQAPRDLIFRTQFLDHPSPSSAWMEPRYKEAANHCALLQEHAQ
RCYVRGLFRSLQQAQSVTSQDLLTAVDAC

Primary citation

Structure of the lysosomal KICSTOR-GATOR1-SAMTOR nutrient-sensing supercomplex. Lupton, C.J., Bayly-Jones, C., Dong, S. et al. Cell (2026) 189:1185-1200.e28. DOI 10.1016/j.cell.2025.12.005 · PubMed

Other PDB entries of the same protein (UniProt Q8WTW4 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9V0J 2.97 Å, Cryo-EM structure of GATOR1-KICSTOR complex
8FW5 3.08 Å, Chimeric HsGATOR1-SpGtr-SpLam complex
9O5A 3.2 Å, The KICSTOR-GATOR1 complex
7T3B 3.9 Å, GATOR1-RAG-RAGULATOR - GAP Complex
6CES 4.0 Å, Cryo-EM structure of GATOR1-RAG
7T3A 4.0 Å, GATOR1-RAG-RAGULATOR - Inhibitory Complex
7T3C 4.0 Å, GATOR1-RAG-RAGULATOR - Dual Complex
6CET 4.4 Å, Cryo-EM structure of GATOR1
9O5E 5.0 Å, The dimeric KICSTOR-GATOR1 supercomplex

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