9OLO: Vesicle-fusing ATPase

22bin20S complex (NSF-alphaSNAP-2:2 syntaxin-1a:SNAP-25), hydrolyzing, class 19. Determined by electron microscopy at 3.56 Å resolution. Released 6 Aug 2025.

Method: Electron microscopy
Resolution: 3.56 Å
Organisms: Cricetulus griseus, Rattus norvegicus
Chains: 14
Atoms: 40,071
Mol. weight: 748.84 kDa
Ligands: PO4, MG, ATP, ADP
Released: 6 Aug 2025

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

Secondary structure: helices and β-sheets

9OLO contains 261 α-helices and 155 β-strands across 14 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: 32 helices, 17 β-strands

Element	Residues	Length	Sheet
α-helix	225-233	9
α-helix	235-237	3
α-helix	242-248	7
α-helix	251-253	3
β-strand	255-259	5	1
α-helix	267-276	10
α-helix	300-303	4
α-helix	307-315	9
β-strand	323-324	2	2
α-helix	330-332	3
β-strand	340	1	3
α-helix	350-352	3
α-helix	355-358	4
β-strand	367-368	2	2
β-strand	369-373	5	1
α-helix	381-383	3
β-strand	394	1	1
α-helix	400-410	11
α-helix	412-416	5
α-helix	427-432	6
α-helix	438-462	25
α-helix	467-471	5
α-helix	478-486	9
α-helix	498-501	4
α-helix	513-529	17
β-strand	536-539	4	4
β-strand	541-542	2	5
α-helix	549-558	10
β-strand	564-568	5	4
α-helix	570-572	3
α-helix	578-592	15
β-strand	598-602	5	4
α-helix	605-608	4
β-strand	612	1	6
β-strand	617	1	6
α-helix	620-629	10
α-helix	633-634	2
β-strand	640-643	4	4
β-strand	646	1	5
α-helix	649-654	6
β-strand	665-666	2	5
β-strand	670	1	7
α-helix	673-683	11
α-helix	686-688	3
α-helix	690-697	8
α-helix	698-700	3
β-strand	705	1	7
α-helix	707-717	11
α-helix	725-735	11

Chain B: 40 helices, 41 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	8
α-helix	14-17	4
β-strand	22-24	3	8
β-strand	34-40	7	8
β-strand	43-51	9	8
α-helix	55-56	2
β-strand	59-62	4	8
α-helix	64-70	7
β-strand	77-82	6	8
β-strand	91	1	9
β-strand	92-99	8	10
β-strand	100-101	2	11
α-helix	104-106	3
β-strand	112-113	2	12
α-helix	114-124	11
β-strand	128-131	4	9
β-strand	135-139	5	11
β-strand	144-148	5	11
β-strand	149-154	6	10
β-strand	171	1	10
β-strand	174-177	4	9
β-strand	184-187	4	10
α-helix	188	1
β-strand	195-196	2	12
β-strand	200	1	10
α-helix	201-203	3
β-strand	209	1	13
α-helix	215-217	3
α-helix	230-233	4
α-helix	236-238	3
α-helix	242-248	7
β-strand	256-259	4	14
α-helix	267-276	10
β-strand	277	1	13
α-helix	279-282	4
β-strand	283-285	3	14
β-strand	294	1	15
α-helix	298-303	6
α-helix	304-306	3
α-helix	307-315	9
α-helix	320-322	3
β-strand	323-328	6	14
α-helix	330-332	3
α-helix	347-358	12
β-strand	367-372	6	14
α-helix	381-383	3
β-strand	385	1	16
β-strand	388	1	16
β-strand	391-394	4	14
α-helix	400-416	17
β-strand	420	1	17
α-helix	426-432	7
α-helix	438-454	17
α-helix	468-471	4
β-strand	475	1	17
α-helix	477-486	10
α-helix	499-502	4
α-helix	513-529	17
β-strand	538-542	5	18
α-helix	549-558	10
β-strand	565-568	4	18
α-helix	570-572	3
β-strand	576	1	3
α-helix	578-591	14
α-helix	592-594	3
β-strand	599-602	4	18
α-helix	605-608	4
β-strand	612	1	19
β-strand	617	1	19
α-helix	620-629	10
α-helix	633-634	2
β-strand	642-646	5	18
α-helix	649-654	6
β-strand	664-666	3	18
β-strand	670	1	20
α-helix	673-683	11
α-helix	688-698	11
β-strand	705	1	20
α-helix	707-717	11
α-helix	722-724	3
α-helix	725-735	11

Chain C: 30 helices, 21 β-strands

Element	Residues	Length	Sheet
β-strand	209	1	21
α-helix	215-218	4
α-helix	224-235	12
α-helix	236-238	3
α-helix	242-248	7
β-strand	256-259	4	22
α-helix	266-276	11
β-strand	277	1	21
β-strand	283-285	3	22
β-strand	294	1	23
α-helix	298-303	6
α-helix	307-316	10
α-helix	320-322	3
β-strand	323-327	5	22
α-helix	330-332	3
α-helix	346-358	13
β-strand	367-373	7	22
β-strand	391-394	4	22
α-helix	400-410	11
α-helix	413-416	4
β-strand	420	1	24
α-helix	426-432	7
α-helix	438-456	19
β-strand	458	1	25
β-strand	464	1	25
α-helix	467-471	5
β-strand	475	1	24
α-helix	477-486	10
α-helix	498-501	4
α-helix	513-529	17
β-strand	537-542	6	26
α-helix	550-558	9
β-strand	565-568	4	26
α-helix	570-572	3
α-helix	578-592	15
β-strand	599-602	4	26
α-helix	605-608	4
β-strand	612	1	27
β-strand	617	1	27
α-helix	621-627	7
α-helix	632-634	3
β-strand	641-646	6	26
α-helix	649-654	6
β-strand	664-666	3	26
α-helix	668-669	2
β-strand	670	1	28
α-helix	673-683	11
α-helix	689-698	10
β-strand	705	1	28
α-helix	707-716	10
α-helix	725-736	12

Chain D: 33 helices, 40 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	29
α-helix	14-17	4
β-strand	22-24	3	29
β-strand	34-38	5	29
β-strand	44-51	8	29
α-helix	55-56	2
β-strand	59-62	4	29
α-helix	64-70	7
β-strand	77-83	7	29
β-strand	91	1	30
β-strand	92-101	10	31
α-helix	104-106	3
β-strand	112-113	2	32
α-helix	114-125	12
β-strand	128-131	4	30
β-strand	135-140	6	31
β-strand	143-152	10	31
β-strand	153	1	33
β-strand	154	1	31
β-strand	171	1	33
β-strand	174-177	4	30
β-strand	182-187	6	31
α-helix	188	1
β-strand	195-196	2	32
β-strand	209	1	34
α-helix	225-233	9
α-helix	244-248	5
α-helix	251-253	3
β-strand	256-259	4	35
α-helix	266-276	11
β-strand	277	1	34
α-helix	281-282	2
β-strand	283-285	3	35
α-helix	298-303	6
α-helix	308-315	8
β-strand	323-327	5	35
α-helix	330-333	4
α-helix	346-348	3
α-helix	350-358	9
β-strand	367-373	7	35
α-helix	381-383	3
β-strand	386	1	36
β-strand	388	1	36
β-strand	391-392	2	35
α-helix	400-415	16
β-strand	420	1	37
α-helix	426-432	7
α-helix	438-454	17
β-strand	457-459	3	38
β-strand	463-465	3	38
β-strand	475	1	37
α-helix	477-486	10
α-helix	496-502	7
α-helix	513-527	15
β-strand	536-542	7	39
α-helix	550-559	10
β-strand	564-568	5	39
α-helix	570-572	3
α-helix	578-590	13
β-strand	597-602	6	39
α-helix	605-608	4
β-strand	611-612	2	40
β-strand	617-618	2	40
α-helix	620-629	10
β-strand	639-645	7	39
α-helix	649-654	6
β-strand	665-666	2	39
β-strand	670	1	41
α-helix	673-683	11
α-helix	688-698	11
β-strand	705	1	41
α-helix	710-717	8
α-helix	726-735	10

Chain E: 28 helices, 18 β-strands

Element	Residues	Length	Sheet
α-helix	225-233	9
α-helix	242-248	7
β-strand	256-259	4	42
α-helix	266-275	10
β-strand	283-285	3	42
α-helix	287-289	3
α-helix	297-303	7
α-helix	307-315	9
β-strand	323-328	6	42
α-helix	346-358	13
β-strand	367-373	7	42
α-helix	376-378	3
α-helix	381-384	4
β-strand	392-394	3	42
α-helix	401-409	9
α-helix	412-416	5
β-strand	420	1	43
α-helix	426-432	7
α-helix	438-455	18
β-strand	457-458	2	44
β-strand	464-465	2	44
β-strand	475	1	43
α-helix	477-486	10
α-helix	496-502	7
α-helix	513-529	17
β-strand	536-541	6	45
α-helix	549-559	11
β-strand	566-568	3	45
α-helix	570-572	3
α-helix	578-592	15
β-strand	600-603	4	45
α-helix	605-608	4
β-strand	612	1	46
β-strand	617	1	46
α-helix	620-629	10
α-helix	632-634	3
β-strand	640-645	6	45
α-helix	649-654	6
β-strand	664-665	2	45
β-strand	670	1	47
α-helix	673-683	11
α-helix	689-697	9
β-strand	705	1	47
α-helix	707-717	11
α-helix	722-724	3
α-helix	725-736	12

Chain F: 28 helices, 16 β-strands

Element	Residues	Length	Sheet
α-helix	215-218	4
α-helix	226-238	13
α-helix	242-248	7
α-helix	250-253	4
β-strand	256-259	4	48
α-helix	266-276	11
β-strand	283-285	3	48
α-helix	297-316	20
β-strand	324-328	5	48
α-helix	330-333	4
α-helix	346-359	14
β-strand	368-373	6	48
α-helix	376-378	3
α-helix	381-384	4
β-strand	391-394	4	48
α-helix	400-415	16
β-strand	420	1	49
α-helix	426-432	7
α-helix	438-458	21
β-strand	475	1	49
α-helix	477-486	10
α-helix	488-490	3
α-helix	495-502	8
α-helix	513-529	17
β-strand	536-542	7	50
α-helix	549-559	11
β-strand	564-568	5	50
α-helix	570-572	3
α-helix	578-592	15
β-strand	597-602	6	50
α-helix	605-608	4
β-strand	612	1	51
β-strand	617	1	51
α-helix	620-629	10
β-strand	639-646	8	50
α-helix	649-654	6
β-strand	665-666	2	50
α-helix	668-669	2
β-strand	670	1	52
α-helix	673-683	11
α-helix	688-698	11
β-strand	705	1	52
α-helix	707-717	11
α-helix	726-735	10

Chain G: 1 helix, 0 β-strands

Element	Residues	Length	Sheet
α-helix	197-256	60

Chain H: 3 helices, 2 β-strands

Element	Residues	Length	Sheet
β-strand	185	1	15
β-strand	187	1	23
α-helix	202-206	5
α-helix	207-213	7
α-helix	214-257	44

6 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
Vesicle-fusing ATPase	A, B, C, D, E, F	protein	747	Cricetulus griseus	P18708 (AlphaFold model)
Syntaxin-1A	G, H	protein	267	Rattus norvegicus	P32851 (AlphaFold model)
Synaptosomal-associated protein 25	I, J	protein	222	Rattus norvegicus	P60881 (AlphaFold model)
Alpha-soluble NSF attachment protein	K, L, M, N	protein	296	Rattus norvegicus	P54921 (AlphaFold model)

Sequence of entity 1 (A, B, C, D, E, F), FASTA

>9OLO_1 Vesicle-fusing ATPase (chains A, B, C, D, E, F)
GANMAGRSMQAARCPTDELSLSNCAVVSEKDYQSGQHVIVRTSPNHKYIFTLRTHPSVVP
GSVAFSLPQRKWAGLSIGQEIEVALYSFDKAKQCIGTMTIEIDFLQKKNIDSNPYDTDKM
AAEFIQQFNNQAFSVGQQLVFSFNDKLFGLLVKDIEAMDPSILKGEPASGKRQKIEVGLV
VGNSQVAFEKAENSSLNLIGKAKTKENRQSIINPDWNFEKMGIGGLDKEFSDIFRRAFAS
RVFPPEIVEQMGCKHVKGILLYGPPGCGKTLLARQIGKMLNAREPKVVNGPEILNKYVGE
SEANIRKLFADAEEEQRRLGANSGLHIIIFDEIDAICKQRGSMAGSTGVHDTVVNQLLSK
IDGVEQLNNILVIGMTNRPDLIDEALLRPGRLEVKMEIGLPDEKGRLQILHIHTARMRGH
QLLSADVDIKELAVETKNFSGAELEGLVRAAQSTAMNRHIKASTKVEVDMEKAESLQVTR
GDFLASLENDIKPAFGTNQEDYASYIMNGIIKWGDPVTRVLDDGELLVQQTKNSDRTPLV
SVLLEGPPHSGKTALAAKIAEESNFPFIKICSPDKMIGFSETAKCQAMKKIFDDAYKSQL
SCVVVDDIERLLDYVPIGPRFSNLVLQALLVLLKKAPPQGRKLLIIGTTSRKDVLQEMEM
LNAFSTTIHVPNIATGEQLLEALELLGNFKDKERTTIAQQVKGKKVWIGIKKLLMLIEMS
LQMDPEYRVRKFLALLREEGASPLDFD

Sequence of entity 2 (G, H), FASTA

>9OLO_2 Syntaxin-1A (chains G, H)
MKDRTQELRTAKDSDDDDDVTVTVDRDRFMDEFFEQVEEIRGFIDKIAENVEEVKRKHSA
ILASPNPDEKTKEELEELMSDIKKTANKVRSKLKSIEQSIEQEEGLNRSSADLRIRKTQH
STLSRKFVEVMSEYNATQSDYRERCKGRIQRQLEITGRTTTSEELEDMLESGNPAIFASG
IIMDSSISKQALSEIETRHSEIIKLENSIRELHDMFMDMAMLVESQGEMIDRIEYNVEHA
VDYVERAVSDTKKAVKYQSKARRKKIM

Sequence of entity 3 (I, J), FASTA

>9OLO_3 Synaptosomal-associated protein 25 (chains I, J)
MGSSHHHHHHSQDPNSMAEDADMRNELEEMQRRADQLADESLESTRRMLQLVEESKDAGI
RTLVMLDEQGEQLERIEEGMDQINKDMKEAEKNLTDLGKFAGLAVAPANKLKSSDAYKKA
WGNNQDGVVASQPARVVDEREQMAISGGFIRRVTNDARENEMDENLEQVSGIIGNLRHMA
LDMGNEIDTQNRQIDRIMEKADSNKTRIDEANQRATKMLGSG

Sequence of entity 4 (K, L, M, N), FASTA

>9OLO_4 Alpha-soluble NSF attachment protein (chains K, L, M, N)
GMDTSGKQAEAMALLAEAERKVKNSQSFFSGLFGGSSKIEEACEIYARAANMFKMAKNWS
AAGNAFCQAAQLHLQLQSKHDAATCFVDAGNAFKKADPQEAINCLMRAIEIYTDMGRFTI
AAKHHISIAEIYETELVDVEKAIAHYEQSADYYKGEESNSSANKCLLKVAGYAAQLEQYQ
KAIDIYEQVGTSAMDSPLLKYSAKDYFFKAALCHFCIDMLNAKLAVQKYEELFPAFSDSR
ECKLMKKLLEAHEEQNVDSYTESVKEYDSISRLDQWLTTMLLRIKKTIQGDEEDLR

Ligands and cofactors

ID	Name	Formula	Copies
PO4	Phosphate ion	O4 P	3
MG	Magnesium ion	Mg	10
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	7
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	5

Primary citation

Structural remodeling of target-SNARE protein complexes by NSF enables synaptic transmission. White, K.I., Khan, Y.A., Qiu, K. et al. Nat Commun (2025) 16:8371-8371. DOI 10.1038/s41467-025-62764-0 · PubMed

Other PDB entries of the same protein (UniProt P18708 (AlphaFold model), which also has an AlphaFold model), best resolution first:

1D2N 1.75 Å, D2 domain of N-ethylmaleimide-sensitive fusion protein
1NSF 1.9 Å, D2 hexamerization domain of N-ethylmaleimide sensitive factor (NSF)
1QCS 1.9 Å, N-terminal domain of N-ethylmaleimide sensitive factor (NSF)
1QDN 2.3 Å, Amino terminal domain of the N-ethylmaleimide sensitive fusion protein (NSF)
9OJR 2.95 Å, 21bin20S complex (NSF-alphaSNAP-2:1 syntaxin-1a:SNAP-25), non-hydrolyzing, class 3
9OJU 2.97 Å, 21bin20S complex (NSF-alphaSNAP-2:1 syntaxin-1a:SNAP-25), non-hydrolyzing, class 4
9PFF 3.09 Å, Min22bin20S complex (NSF-alphaSNAP-2:2 syntaxin-1a H3:SNAP-25 SN1), non-hydrolyzing,…
9PFC 3.15 Å, NSF, substrate free, hydrolyzing, class 26
9OK5 3.29 Å, 22bin20S complex (NSF-alphaSNAP-2:2 syntaxin-1a:SNAP-25), hydrolyzing, class 16
9OJZ 3.39 Å, 21bin20S complex (NSF-alphaSNAP-2:1 syntaxin-1a:SNAP-25), non-hydrolyzing, class 5
9PB9 3.45 Å, 21bin20S complex (NSF-alphaSNAP-2:1 syntaxin-1a:SNAP-25), non-hydrolyzing, class 8
9PBA 3.47 Å, 21bin20S complex (NSF-alphaSNAP-2:1 syntaxin-1a:SNAP-25), non-hydrolyzing, class 9

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