9PFC: NSF, substrate free, hydrolyzing, class 26
NSF, substrate free, hydrolyzing, class 26. Determined by electron microscopy at 3.15 Å resolution. Released 8 Oct 2025.
- Method
- Electron microscopy
- Resolution
- 3.15 Å
- Organism
- Cricetulus griseus
- Chains
- 6
- Atoms
- 29,491
- Mol. weight
- 503.54 kDa
- Ligands
- PO4, MG, ADP, ATP
- Released
- 8 Oct 2025
Explore 9PFC in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9PFC contains 189 α-helices and 143 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: 36 helices, 36 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 4-10 | 7 | 1 |
| α-helix | 14-19 | 6 | |
| β-strand | 22-24 | 3 | 1 |
| β-strand | 34-40 | 7 | 1 |
| β-strand | 43-51 | 9 | 1 |
| α-helix | 56 | 1 | |
| β-strand | 59-62 | 4 | 1 |
| α-helix | 64-70 | 7 | |
| β-strand | 77-83 | 7 | 1 |
| α-helix | 87-90 | 4 | |
| β-strand | 91 | 1 | 2 |
| β-strand | 92-101 | 10 | 3 |
| α-helix | 104-106 | 3 | |
| β-strand | 111-113 | 3 | 4 |
| α-helix | 114-125 | 12 | |
| β-strand | 128-131 | 4 | 2 |
| β-strand | 135-140 | 6 | 3 |
| β-strand | 143-154 | 12 | 3 |
| α-helix | 155-158 | 4 | |
| β-strand | 171 | 1 | 3 |
| β-strand | 174-177 | 4 | 2 |
| β-strand | 182-187 | 6 | 3 |
| α-helix | 188 | 1 | |
| β-strand | 194-196 | 3 | 4 |
| α-helix | 215-218 | 4 | |
| α-helix | 225-234 | 10 | |
| β-strand | 256-259 | 4 | 5 |
| α-helix | 266-276 | 11 | |
| α-helix | 282 | 1 | |
| β-strand | 283 | 1 | 5 |
| α-helix | 284 | 1 | |
| α-helix | 287-290 | 4 | |
| α-helix | 298-315 | 18 | |
| α-helix | 320-322 | 3 | |
| β-strand | 323-327 | 5 | 5 |
| α-helix | 330-333 | 4 | |
| α-helix | 348-358 | 11 | |
| β-strand | 367-373 | 7 | 5 |
| α-helix | 376-378 | 3 | |
| β-strand | 385 | 1 | 6 |
| β-strand | 388 | 1 | 6 |
| β-strand | 392-394 | 3 | 5 |
| α-helix | 400-410 | 11 | |
| α-helix | 413-417 | 5 | |
| β-strand | 420 | 1 | 7 |
| α-helix | 426-432 | 7 | |
| α-helix | 438-461 | 24 | |
| α-helix | 467-472 | 6 | |
| β-strand | 476 | 1 | 7 |
| α-helix | 479-486 | 8 | |
| β-strand | 495 | 1 | 8 |
| α-helix | 512-525 | 14 | |
| β-strand | 536-542 | 7 | 9 |
| α-helix | 550-560 | 11 | |
| β-strand | 564 | 1 | 8 |
| β-strand | 566-568 | 3 | 9 |
| α-helix | 570-572 | 3 | |
| α-helix | 578-592 | 15 | |
| β-strand | 600-603 | 4 | 9 |
| α-helix | 605-608 | 4 | |
| β-strand | 612 | 1 | 10 |
| β-strand | 617 | 1 | 10 |
| α-helix | 620-629 | 10 | |
| β-strand | 640-646 | 7 | 9 |
| α-helix | 650-654 | 5 | |
| β-strand | 664-666 | 3 | 9 |
| β-strand | 670 | 1 | 11 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-697 | 10 | |
| β-strand | 705 | 1 | 11 |
| α-helix | 707-717 | 11 | |
| α-helix | 726-735 | 10 | |
Chain B: 36 helices, 35 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 4-10 | 7 | 12 |
| α-helix | 14-19 | 6 | |
| β-strand | 22-24 | 3 | 12 |
| β-strand | 34-40 | 7 | 12 |
| β-strand | 43-51 | 9 | 12 |
| α-helix | 56 | 1 | |
| β-strand | 59-62 | 4 | 12 |
| α-helix | 64-70 | 7 | |
| β-strand | 77-83 | 7 | 12 |
| α-helix | 87-90 | 4 | |
| β-strand | 91 | 1 | 13 |
| β-strand | 92-101 | 10 | 14 |
| α-helix | 104-106 | 3 | |
| β-strand | 111-113 | 3 | 15 |
| α-helix | 114-125 | 12 | |
| β-strand | 129-131 | 3 | 13 |
| β-strand | 135-140 | 6 | 14 |
| β-strand | 143-154 | 12 | 14 |
| α-helix | 155-158 | 4 | |
| β-strand | 171 | 1 | 14 |
| β-strand | 174-176 | 3 | 13 |
| β-strand | 182-187 | 6 | 14 |
| α-helix | 188 | 1 | |
| β-strand | 194-196 | 3 | 15 |
| α-helix | 215-218 | 4 | |
| α-helix | 225-230 | 6 | |
| α-helix | 231-235 | 5 | |
| α-helix | 242-248 | 7 | |
| β-strand | 255-259 | 5 | 16 |
| α-helix | 266-276 | 11 | |
| α-helix | 280-282 | 3 | |
| α-helix | 287-290 | 4 | |
| α-helix | 299-306 | 8 | |
| α-helix | 307-309 | 3 | |
| α-helix | 310-316 | 7 | |
| β-strand | 323-327 | 5 | 16 |
| α-helix | 348-358 | 11 | |
| β-strand | 367-373 | 7 | 16 |
| β-strand | 386 | 1 | 17 |
| β-strand | 388 | 1 | 17 |
| β-strand | 392-394 | 3 | 16 |
| α-helix | 400-410 | 11 | |
| α-helix | 413-416 | 4 | |
| β-strand | 420 | 1 | 18 |
| α-helix | 426-432 | 7 | |
| α-helix | 444-454 | 11 | |
| β-strand | 458-459 | 2 | 19 |
| β-strand | 463-464 | 2 | 19 |
| β-strand | 475 | 1 | 18 |
| α-helix | 477-481 | 5 | |
| α-helix | 496-502 | 7 | |
| α-helix | 512-527 | 16 | |
| β-strand | 536-542 | 7 | 20 |
| α-helix | 551-558 | 8 | |
| β-strand | 566-568 | 3 | 20 |
| α-helix | 570-572 | 3 | |
| α-helix | 578-592 | 15 | |
| β-strand | 600-603 | 4 | 20 |
| α-helix | 605-608 | 4 | |
| β-strand | 612 | 1 | 21 |
| β-strand | 617 | 1 | 21 |
| α-helix | 620-629 | 10 | |
| β-strand | 640-646 | 7 | 20 |
| α-helix | 649-654 | 6 | |
| β-strand | 664-666 | 3 | 20 |
| β-strand | 670 | 1 | 22 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-698 | 11 | |
| β-strand | 705 | 1 | 22 |
| α-helix | 707-713 | 7 | |
| α-helix | 725-735 | 11 | |
Chain C: 27 helices, 19 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 226-235 | 10 | |
| α-helix | 242-248 | 7 | |
| β-strand | 256-259 | 4 | 23 |
| α-helix | 267-276 | 10 | |
| β-strand | 283-284 | 2 | 23 |
| α-helix | 287-290 | 4 | |
| α-helix | 298-303 | 6 | |
| α-helix | 307-316 | 10 | |
| β-strand | 323-326 | 4 | 23 |
| β-strand | 340 | 1 | 24 |
| α-helix | 350-358 | 9 | |
| β-strand | 367-373 | 7 | 23 |
| α-helix | 376-378 | 3 | |
| β-strand | 391-394 | 4 | 23 |
| α-helix | 400-410 | 11 | |
| β-strand | 420 | 1 | 25 |
| α-helix | 426-432 | 7 | |
| α-helix | 438-452 | 15 | |
| β-strand | 457-458 | 2 | 26 |
| β-strand | 464-465 | 2 | 26 |
| α-helix | 470-472 | 3 | |
| β-strand | 475 | 1 | 25 |
| α-helix | 477-486 | 10 | |
| α-helix | 496-498 | 3 | |
| α-helix | 513-530 | 18 | |
| β-strand | 536-542 | 7 | 27 |
| α-helix | 549-560 | 12 | |
| β-strand | 566-568 | 3 | 27 |
| α-helix | 570-572 | 3 | |
| α-helix | 578-590 | 13 | |
| α-helix | 591-594 | 4 | |
| β-strand | 600-602 | 3 | 27 |
| α-helix | 605-608 | 4 | |
| β-strand | 612 | 1 | 28 |
| β-strand | 617 | 1 | 28 |
| α-helix | 620-629 | 10 | |
| α-helix | 633-634 | 2 | |
| β-strand | 640-646 | 7 | 27 |
| α-helix | 649-654 | 6 | |
| β-strand | 664-666 | 3 | 27 |
| β-strand | 670 | 1 | 29 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-698 | 11 | |
| β-strand | 705 | 1 | 29 |
| α-helix | 707-716 | 10 | |
| α-helix | 725-736 | 12 | |
Chain D: 28 helices, 23 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 216-218 | 3 | |
| α-helix | 225-230 | 6 | |
| α-helix | 231-235 | 5 | |
| α-helix | 242-248 | 7 | |
| β-strand | 255-259 | 5 | 30 |
| α-helix | 266-276 | 11 | |
| α-helix | 280-283 | 4 | |
| β-strand | 285-286 | 2 | 30 |
| α-helix | 287-290 | 4 | |
| α-helix | 297-316 | 20 | |
| β-strand | 323-328 | 6 | 30 |
| α-helix | 348-359 | 12 | |
| β-strand | 367-373 | 7 | 30 |
| α-helix | 376-378 | 3 | |
| α-helix | 381-383 | 3 | |
| β-strand | 391-394 | 4 | 30 |
| α-helix | 400-406 | 7 | |
| β-strand | 420 | 1 | 31 |
| α-helix | 426-429 | 4 | |
| α-helix | 441-452 | 12 | |
| α-helix | 453-456 | 4 | |
| β-strand | 457-458 | 2 | 32 |
| β-strand | 464-465 | 2 | 32 |
| β-strand | 475 | 1 | 31 |
| α-helix | 477-486 | 10 | |
| β-strand | 494 | 1 | 33 |
| α-helix | 513-530 | 18 | |
| β-strand | 536-542 | 7 | 34 |
| α-helix | 550-558 | 9 | |
| β-strand | 565 | 1 | 33 |
| β-strand | 566-568 | 3 | 34 |
| α-helix | 570-572 | 3 | |
| β-strand | 576 | 1 | 24 |
| α-helix | 578-590 | 13 | |
| α-helix | 591-594 | 4 | |
| β-strand | 597 | 1 | 35 |
| β-strand | 600-602 | 3 | 34 |
| α-helix | 605-608 | 4 | |
| β-strand | 612 | 1 | 36 |
| β-strand | 617 | 1 | 36 |
| α-helix | 620-629 | 10 | |
| β-strand | 639 | 1 | 35 |
| β-strand | 640-646 | 7 | 34 |
| α-helix | 649-652 | 4 | |
| β-strand | 664-666 | 3 | 34 |
| β-strand | 670 | 1 | 37 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-698 | 11 | |
| β-strand | 705 | 1 | 37 |
| α-helix | 713-716 | 4 | |
| α-helix | 725-736 | 12 | |
Chain E: 34 helices, 16 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 225-234 | 10 | |
| α-helix | 236-238 | 3 | |
| α-helix | 242-248 | 7 | |
| α-helix | 251-253 | 3 | |
| β-strand | 256-259 | 4 | 38 |
| α-helix | 266-276 | 11 | |
| α-helix | 281-282 | 2 | |
| β-strand | 283-284 | 2 | 38 |
| α-helix | 287-291 | 5 | |
| α-helix | 297-305 | 9 | |
| α-helix | 307-315 | 9 | |
| α-helix | 320-322 | 3 | |
| β-strand | 323-326 | 4 | 38 |
| α-helix | 330-333 | 4 | |
| α-helix | 347-358 | 12 | |
| β-strand | 367-373 | 7 | 38 |
| α-helix | 376-378 | 3 | |
| α-helix | 381-384 | 4 | |
| β-strand | 391-394 | 4 | 38 |
| α-helix | 402-410 | 9 | |
| α-helix | 413-416 | 4 | |
| β-strand | 420 | 1 | 39 |
| α-helix | 426-432 | 7 | |
| α-helix | 438-459 | 22 | |
| β-strand | 475 | 1 | 39 |
| α-helix | 477-486 | 10 | |
| α-helix | 512-514 | 3 | |
| α-helix | 522-530 | 9 | |
| β-strand | 536-542 | 7 | 40 |
| α-helix | 549-558 | 10 | |
| β-strand | 566-568 | 3 | 40 |
| α-helix | 570-572 | 3 | |
| α-helix | 580-590 | 11 | |
| α-helix | 591-593 | 3 | |
| β-strand | 600-602 | 3 | 40 |
| α-helix | 605-609 | 5 | |
| β-strand | 612 | 1 | 41 |
| β-strand | 617 | 1 | 41 |
| α-helix | 620-629 | 10 | |
| β-strand | 640-646 | 7 | 40 |
| α-helix | 649-652 | 4 | |
| α-helix | 653-655 | 3 | |
| β-strand | 664-666 | 3 | 40 |
| β-strand | 670 | 1 | 42 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-698 | 11 | |
| β-strand | 705 | 1 | 42 |
| α-helix | 707-717 | 11 | |
| α-helix | 722-724 | 3 | |
| α-helix | 725-736 | 12 | |
Chain F: 28 helices, 14 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 216-218 | 3 | |
| α-helix | 224-233 | 10 | |
| β-strand | 256-259 | 4 | 43 |
| α-helix | 267-276 | 10 | |
| α-helix | 280-282 | 3 | |
| β-strand | 283-286 | 4 | 43 |
| α-helix | 298-316 | 19 | |
| β-strand | 323-328 | 6 | 43 |
| α-helix | 347-358 | 12 | |
| β-strand | 367-373 | 7 | 43 |
| α-helix | 376-378 | 3 | |
| α-helix | 382-384 | 3 | |
| β-strand | 391-394 | 4 | 43 |
| α-helix | 400-410 | 11 | |
| α-helix | 412-416 | 5 | |
| α-helix | 426-431 | 6 | |
| α-helix | 438-456 | 19 | |
| α-helix | 477-486 | 10 | |
| α-helix | 498-500 | 3 | |
| α-helix | 512-527 | 16 | |
| β-strand | 537-542 | 6 | 44 |
| α-helix | 549-558 | 10 | |
| β-strand | 564-568 | 5 | 44 |
| α-helix | 570-572 | 3 | |
| α-helix | 578-592 | 15 | |
| β-strand | 597-602 | 6 | 44 |
| α-helix | 605-608 | 4 | |
| β-strand | 612 | 1 | 45 |
| β-strand | 617 | 1 | 45 |
| α-helix | 620-629 | 10 | |
| α-helix | 632-634 | 3 | |
| β-strand | 639-645 | 7 | 44 |
| α-helix | 649-654 | 6 | |
| α-helix | 658-660 | 3 | |
| β-strand | 663-666 | 4 | 44 |
| β-strand | 670-672 | 3 | 46 |
| α-helix | 673-683 | 11 | |
| α-helix | 688-698 | 11 | |
| β-strand | 702-705 | 4 | 46 |
| α-helix | 708-718 | 11 | |
| α-helix | 722-724 | 3 | |
| α-helix | 727-736 | 10 | |
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) |
Sequence of entity 1 (A, B, C, D, E, F), FASTA
>9PFC_1 Vesicle-fusing ATPase (chains A, B, C, D, E, F)
GANMAGRSMQAARCPTDELSLSNCAVVSEKDYQSGQHVIVRTSPNHKYIFTLRTHPSVVP
GSVAFSLPQRKWAGLSIGQEIEVALYSFDKAKQCIGTMTIEIDFLQKKNIDSNPYDTDKM
AAEFIQQFNNQAFSVGQQLVFSFNDKLFGLLVKDIEAMDPSILKGEPASGKRQKIEVGLV
VGNSQVAFEKAENSSLNLIGKAKTKENRQSIINPDWNFEKMGIGGLDKEFSDIFRRAFAS
RVFPPEIVEQMGCKHVKGILLYGPPGCGKTLLARQIGKMLNAREPKVVNGPEILNKYVGE
SEANIRKLFADAEEEQRRLGANSGLHIIIFDEIDAICKQRGSMAGSTGVHDTVVNQLLSK
IDGVEQLNNILVIGMTNRPDLIDEALLRPGRLEVKMEIGLPDEKGRLQILHIHTARMRGH
QLLSADVDIKELAVETKNFSGAELEGLVRAAQSTAMNRHIKASTKVEVDMEKAESLQVTR
GDFLASLENDIKPAFGTNQEDYASYIMNGIIKWGDPVTRVLDDGELLVQQTKNSDRTPLV
SVLLEGPPHSGKTALAAKIAEESNFPFIKICSPDKMIGFSETAKCQAMKKIFDDAYKSQL
SCVVVDDIERLLDYVPIGPRFSNLVLQALLVLLKKAPPQGRKLLIIGTTSRKDVLQEMEM
LNAFSTTIHVPNIATGEQLLEALELLGNFKDKERTTIAQQVKGKKVWIGIKKLLMLIEMS
LQMDPEYRVRKFLALLREEGASPLDFD
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| PO4 | Phosphate ion | O4 P | 4 |
| MG | Magnesium ion | Mg | 7 |
| ADP | Adenosine-5'-diphosphate | C10 H15 N5 O10 P2 | 5 |
| ATP | Adenosine-5'-triphosphate | C10 H16 N5 O13 P3 | 7 |
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,…
- 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
- 9OLJ 3.52 Å, 22bin20S complex (NSF-alphaSNAP-2:2 syntaxin-1a:SNAP-25), hydrolyzing, class 18
Browse structure collections
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