PDBsum entry 1d2n

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Hexamerization domain PDB id
Protein chain
246 a.a. *
Waters ×271
* Residue conservation analysis
PDB id:
Name: Hexamerization domain
Title: D2 domain of n-ethylmaleimide-sensitive fusion protein
Structure: N-ethylmaleimide-sensitive fusion protein. Chain: a. Fragment: d2. Engineered: yes
Source: Cricetulus griseus. Chinese hamster. Organism_taxid: 10029. Organ: ovary. Cellular_location: cytoplasm. Expressed in: escherichia coli. Expression_system_taxid: 562.
Biol. unit: Hexamer (from PDB file)
1.75Å     R-factor:   0.199     R-free:   0.223
Authors: C.U.Lenzen,D.Steinmann,S.W.Whiteheart,W.I.Weis
Key ref:
C.U.Lenzen et al. (1998). Crystal structure of the hexamerization domain of N-ethylmaleimide-sensitive fusion protein. Cell, 94, 525-536. PubMed id: 9727495 DOI: 10.1016/S0092-8674(00)81593-7
30-Jun-98     Release date:   14-Oct-98    
Go to PROCHECK summary

Protein chain
Pfam   ArchSchema ?
P18708  (NSF_CRIGR) -  Vesicle-fusing ATPase
744 a.a.
246 a.a.
Key:    PfamA domain  Secondary structure  CATH domain

 Enzyme reactions 
   Enzyme class: E.C.  - Vesicle-fusing ATPase.
[IntEnz]   [ExPASy]   [KEGG]   [BRENDA]
      Reaction: ATP + H2O = ADP + phosphate
+ H(2)O
Bound ligand (Het Group name = ANP)
matches with 81.25% similarity
+ phosphate
Molecule diagrams generated from .mol files obtained from the KEGG ftp site
 Gene Ontology (GO) functional annotation 
  GO annot!
  Biochemical function     nucleotide binding     3 terms  


DOI no: 10.1016/S0092-8674(00)81593-7 Cell 94:525-536 (1998)
PubMed id: 9727495  
Crystal structure of the hexamerization domain of N-ethylmaleimide-sensitive fusion protein.
C.U.Lenzen, D.Steinmann, S.W.Whiteheart, W.I.Weis.
N-ethylmaleimide-sensitive fusion protein (NSF) is a cytosolic ATPase required for many intracellular vesicle fusion reactions. NSF consists of an amino-terminal region that interacts with other components of the vesicle trafficking machinery, followed by two homologous ATP-binding cassettes, designated D1 and D2, that possess essential ATPase and hexamerization activities, respectively. The crystal structure of D2 bound to Mg2+-AMPPNP has been determined at 1.75 A resolution. The structure consists of a nucleotide-binding and a helical domain, and it is unexpectedly similar to the first two domains of the clamp-loading subunit delta' of E. coli DNA polymerase III. The structure suggests several regions responsible for coupling of ATP hydrolysis to structural changes in full-length NSF.
  Selected figure(s)  
Figure 1.
Figure 1. Structure of the NSF D2 Protomer(A) Topology of D2. Helices are shown as cylinders; strands are shown as as arrows. Residue numbers at the beginning and end of the principal elements of secondary structure are indicated, as well as the locations of the P loop and the DExx box (DDIE in D2). Helices and strands are numbered consecutively in sequence.(B) Stereo C[α] trace of D2. Every tenth C[α] is shown as a small sphere and numbered. The bound AMPPNP is shown with thickened black bonds.(C) Ribbon diagram of D2, with AMPPNP shown in a ball-and-stick representation. To aid in following the path of the backbone, the ribbon is white at the N terminus and becomes progressively darker moving toward the C terminus. The P loop and DExx box are green and red, respectively.(D) Ribbon diagram of the first two domains of E. coli DNA polymerase III δ′ ([22]), shown in the same orientation and color scheme as (C), except for a zinc-binding insert unique to δ′ (α3), which is shown in white. (B)–(D), as well as Figure 2B and Figure 3, were prepared with MOLSCRIPT ( [28]).
Figure 2.
Figure 2. Nucleotide Binding by D2(A) Stereo view of the refined 2F[o]−F[c] electron density map within 2.4 Å of AMPPNP, contoured at 1.5 σ. The refined model is shown with AMPPNP and Mg^2PLUSPUSSIGN in black, and protein and water molecules in white. The figure was prepared with BOBSCRIPT ([18]).(B) Stereo view of nucleotide-binding site. AMPPNP is shown with black bonds. White, light gray, dark gray, and black spheres denote carbon, nitrogen, oxygen, and phosphorus atoms, respectively. Mg^2PLUSPUSSIGN is shown as a larger black sphere. Water molecules are shown as single oxygen atoms. Hydrogen bonds are shown as thin dashed lines; Mg^2PLUSPUSSIGN coordination bonds are shown as thick dashed lines. For clarity, the backbone at position 510 and the water molecule that interacts with the α-phosphate oxygens (see [C]) are not shown.(C) Schematic diagram of the interactions between AMPPNP and D2. Water molecules are indicated by “W.” Hydrogen and Mg^2PLUSPUSSIGN coordination bonds are indicated with dashed lines. Main-chain amide and carbonyl oxygen groups that interact with the ligand are shown emanating from the box surrounding the residue name, and side chain functionalities are shown schematically. Nonpolar van der Waals contacts are indicated by arcs. In (B) and (C), the asterisk at Lys-639 designates that this residue comes from an adjacent protomer in the D2 hexamer. This lysine appears to be only partially occupied, and its interaction with Oγ of AMPPNP is likely replaced by a water molecule in a fraction of the molecules in the crystal (see text).
  The above figures are reprinted by permission from Cell Press: Cell (1998, 94, 525-536) copyright 1998.  
  Figures were selected by an automated process.  

Literature references that cite this PDB file's key reference

  PubMed id Reference
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Structure and mechanism of the hexameric MecA-ClpC molecular machine.
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PDB codes: 2y1q 2y1r 3pxg 3pxi
19887446 C.Zhao, E.A.Matveeva, Q.Ren, and S.W.Whiteheart (2010).
Dissecting the N-ethylmaleimide-sensitive factor: required elements of the N and D1 domains.
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20462489 G.Effantin, T.Ishikawa, G.M.De Donatis, M.R.Maurizi, and A.C.Steven (2010).
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20130681 K.L.Cheung, J.Huen, W.A.Houry, and J.Ortega (2010).
Comparison of the multiple oligomeric structures observed for the Rvb1 and Rvb2 proteins.
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20113510 R.McNally, G.D.Bowman, E.R.Goedken, M.O'Donnell, and J.Kuriyan (2010).
Analysis of the role of PCNA-DNA contacts during clamp loading.
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PDB code: 3k4x
20962217 W.H.Chou, D.Wang, T.McMahon, Z.H.Qi, M.Song, C.Zhang, K.M.Shokat, and R.O.Messing (2010).
GABAA receptor trafficking is regulated by protein kinase C(epsilon) and the N-ethylmaleimide-sensitive factor.
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19278657 M.J.Landsberg, P.R.Vajjhala, R.Rothnagel, A.L.Munn, and B.Hankamer (2009).
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18329872 E.J.Enemark, and L.Joshua-Tor (2008).
On helicases and other motor proteins.
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18332143 L.C.Briggs, G.S.Baldwin, N.Miyata, H.Kondo, X.Zhang, and P.S.Freemont (2008).
Analysis of nucleotide binding to P97 reveals the properties of a tandem AAA hexameric ATPase.
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18647240 N.D.Thomsen, and J.M.Berger (2008).
Structural frameworks for considering microbial protein- and nucleic acid-dependent motor ATPases.
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17553803 A.A.Horwitz, A.Navon, M.Groll, D.M.Smith, C.Reis, and A.L.Goldberg (2007).
ATP-induced structural transitions in PAN, the proteasome-regulatory ATPase complex in Archaea.
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17397838 C.Zhao, J.T.Slevin, and S.W.Whiteheart (2007).
Cellular functions of NSF: not just SNAPs and SNAREs.
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17242399 J.Schumacher, N.Joly, M.Rappas, D.Bradley, S.R.Wigneshweraraj, X.Zhang, and M.Buck (2007).
Sensor I threonine of the AAA+ ATPase transcriptional activator PspF is involved in coupling nucleotide triphosphate hydrolysis to the restructuring of sigma 54-RNA polymerase.
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17202259 K.Imada, T.Minamino, A.Tahara, and K.Namba (2007).
Structural similarity between the flagellar type III ATPase FliI and F1-ATPase subunits.
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PDB code: 2dpy
17324440 T.D.Ziebarth, C.L.Farr, and L.S.Kaguni (2007).
Modular architecture of the hexameric human mitochondrial DNA helicase.
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17012286 A.F.Neuwald (2006).
Hypothesis: bacterial clamp loader ATPase activation through DNA-dependent repositioning of the catalytic base and of a trans-acting catalytic threonine.
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16955075 C.Indiani, and M.O'Donnell (2006).
The replication clamp-loading machine at work in the three domains of life.
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17081110 C.J.Lowenstein, and H.Tsuda (2006).
N-ethylmaleimide-sensitive factor: a redox sensor in exocytosis.
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16581508 I.G.Woods, D.A.Lyons, M.G.Voas, H.M.Pogoda, and W.S.Talbot (2006).
nsf is essential for organization of myelinated axons in zebrafish.
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16689629 J.P.Erzberger, and J.M.Berger (2006).
Evolutionary relationships and structural mechanisms of AAA+ proteins.
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16829961 J.P.Erzberger, M.L.Mott, and J.M.Berger (2006).
Structural basis for ATP-dependent DnaA assembly and replication-origin remodeling.
  Nat Struct Mol Biol, 13, 676-683.
PDB code: 2hcb
16421093 M.O'Donnell (2006).
Replisome architecture and dynamics in Escherichia coli.
  J Biol Chem, 281, 10653-10656.  
16430918 M.Rappas, J.Schumacher, H.Niwa, M.Buck, and X.Zhang (2006).
Structural basis of the nucleotide driven conformational changes in the AAA+ domain of transcription activator PspF.
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PDB codes: 2c96 2c98 2c99 2c9c
16973614 N.Joly, J.Schumacher, and M.Buck (2006).
Heterogeneous nucleotide occupancy stimulates functionality of phage shock protein F, an AAA+ transcriptional activator.
  J Biol Chem, 281, 34997-35007.  
17060327 P.M.Matias, S.Gorynia, P.Donner, and M.A.Carrondo (2006).
Crystal structure of the human AAA+ protein RuvBL1.
  J Biol Chem, 281, 38918-38929.
PDB code: 2c9o
16877706 T.V.Rotanova, I.Botos, E.E.Melnikov, F.Rasulova, A.Gustchina, M.R.Maurizi, and A.Wlodawer (2006).
Slicing a protease: structural features of the ATP-dependent Lon proteases gleaned from investigations of isolated domains.
  Protein Sci, 15, 1815-1828.  
16061814 J.Shen, D.Gai, A.Patrick, W.B.Greenleaf, and X.S.Chen (2005).
The roles of the residues on the channel beta-hairpin and loop structures of simian virus 40 hexameric helicase.
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15866879 L.J.Crowther, A.Yamagata, L.Craig, J.A.Tainer, and M.S.Donnenberg (2005).
The ATPase activity of BfpD is greatly enhanced by zinc and allosteric interactions with other Bfp proteins.
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15678420 M.Groll, M.Bochtler, H.Brandstetter, T.Clausen, and R.Huber (2005).
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16072036 P.I.Hanson, and S.W.Whiteheart (2005).
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15829969 S.J.Riedl, W.Li, Y.Chao, R.Schwarzenbacher, and Y.Shi (2005).
Structure of the apoptotic protease-activating factor 1 bound to ADP.
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PDB code: 1z6t
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Out-of-plane motions in open sliding clamps: molecular dynamics simulations of eukaryotic and archaeal proliferating cell nuclear antigen.
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Substrate recognition by the AAA+ chaperone ClpB.
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15082529 D.L.Pappas, R.Frisch, and M.Weinreich (2004).
The NAD(+)-dependent Sir2p histone deacetylase is a negative regulator of chromosomal DNA replication.
  Genes Dev, 18, 769-781.  
15201901 G.D.Bowman, M.O'Donnell, and J.Kuriyan (2004).
Structural analysis of a eukaryotic sliding DNA clamp-clamp loader complex.
  Nature, 429, 724-730.
PDB code: 1sxj
15310852 J.A.James, A.K.Aggarwal, R.M.Linden, and C.R.Escalante (2004).
Structure of adeno-associated virus type 2 Rep40-ADP complex: insight into nucleotide recognition and catalysis by superfamily 3 helicases.
  Proc Natl Acad Sci U S A, 101, 12455-12460.
PDB code: 1u0j
15178690 J.M.Tkach, and J.R.Glover (2004).
Amino acid substitutions in the C-terminal AAA+ module of Hsp104 prevent substrate recognition by disrupting oligomerization and cause high temperature inactivation.
  J Biol Chem, 279, 35692-35701.  
14728719 J.Weibezahn, B.Bukau, and A.Mogk (2004).
Unscrambling an egg: protein disaggregation by AAA+ proteins.
  Microb Cell Fact, 3, 1.  
14962378 M.R.Maurizi, and D.Xia (2004).
Protein binding and disruption by Clp/Hsp100 chaperones.
  Structure, 12, 175-183.  
14711675 M.Saloheimo, H.Wang, M.Valkonen, T.Vasara, A.Huuskonen, M.Riikonen, T.Pakula, M.Ward, and M.Penttilä (2004).
Characterization of secretory genes ypt1/yptA and nsf1/nsfA from two filamentous fungi: induction of secretory pathway genes of Trichoderma reesei under secretion stress conditions.
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15031730 P.Laksanalamai, T.A.Whitehead, and F.T.Robb (2004).
Minimal protein-folding systems in hyperthermophilic archaea.
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15556993 S.L.Kazmirski, M.Podobnik, T.F.Weitze, M.O'Donnell, and J.Kuriyan (2004).
Structural analysis of the inactive state of the Escherichia coli DNA polymerase clamp-loader complex.
  Proc Natl Acad Sci U S A, 101, 16750-16755.
PDB codes: 1xxh 1xxi
15210950 T.Hishida, Y.W.Han, S.Fujimoto, H.Iwasaki, and H.Shinagawa (2004).
Direct evidence that a conserved arginine in RuvB AAA+ ATPase acts as an allosteric effector for the ATPase activity of the adjacent subunit in a hexamer.
  Proc Natl Acad Sci U S A, 101, 9573-9577.  
15153108 V.Lake, U.Olsson, R.D.Willows, and M.Hansson (2004).
ATPase activity of magnesium chelatase subunit I is required to maintain subunit D in vivo.
  Eur J Biochem, 271, 2182-2188.  
12949490 B.DeLaBarre, and A.T.Brunger (2003).
Complete structure of p97/valosin-containing protein reveals communication between nucleotide domains.
  Nat Struct Biol, 10, 856-863.
PDB code: 1oz4
12774115 D.Li, R.Zhao, W.Lilyestrom, D.Gai, R.Zhang, J.A.DeCaprio, E.Fanning, A.Jochimiak, G.Szakonyi, and X.S.Chen (2003).
Structure of the replicative helicase of the oncoprotein SV40 large tumour antigen.
  Nature, 423, 512-518.
PDB code: 1n25
14514695 D.Y.Kim, and K.K.Kim (2003).
Crystal structure of ClpX molecular chaperone from Helicobacter pylori.
  J Biol Chem, 278, 50664-50670.
PDB code: 1um8
12622725 F.Hayashi, H.Suzuki, R.Iwase, T.Uzumaki, A.Miyake, J.R.Shen, K.Imada, Y.Furukawa, K.Yonekura, K.Namba, and M.Ishiura (2003).
ATP-induced hexameric ring structure of the cyanobacterial circadian clock protein KaiC.
  Genes Cells, 8, 287-296.  
12906833 J.A.James, C.R.Escalante, M.Yoon-Robarts, T.A.Edwards, R.M.Linden, and A.K.Aggarwal (2003).
Crystal structure of the SF3 helicase from adeno-associated virus type 2.
  Structure, 11, 1025-1035.
PDB code: 1s9h
12948749 J.A.Maupin-Furlow, S.J.Kaczowka, C.J.Reuter, K.Zuobi-Hasona, and M.A.Gil (2003).
Archaeal proteasomes: potential in metabolic engineering.
  Metab Eng, 5, 151-163.  
12941689 J.Furst, R.B.Sutton, J.Chen, A.T.Brunger, and N.Grigorieff (2003).
Electron cryomicroscopy structure of N-ethyl maleimide sensitive factor at 11 A resolution.
  EMBO J, 22, 4365-4374.  
12672453 M.Groll, and R.Huber (2003).
Substrate access and processing by the 20S proteasome core particle.
  Int J Biochem Cell Biol, 35, 606-616.  
12480933 M.J.Davey, C.Indiani, and M.O'Donnell (2003).
Reconstitution of the Mcm2-7p heterohexamer, subunit arrangement, and ATP site architecture.
  J Biol Chem, 278, 4491-4499.  
12807884 Q.Wang, C.Song, X.Yang, and C.C.Li (2003).
D1 ring is stable and nucleotide-independent, whereas D2 ring undergoes major conformational changes during the ATPase cycle of p97-VCP.
  J Biol Chem, 278, 32784-32793.  
12962509 R.C.Bruckner, P.L.Gunyuzlu, and R.L.Stein (2003).
Coupled kinetics of ATP and peptide hydrolysis by Escherichia coli FtsH protease.
  Biochemistry, 42, 10843-10852.  
12727865 S.N.Savvides, H.J.Yeo, M.R.Beck, F.Blaesing, R.Lurz, E.Lanka, R.Buhrdorf, W.Fischer, R.Haas, and G.Waksman (2003).
VirB11 ATPases are dynamic hexameric assemblies: new insights into bacterial type IV secretion.
  EMBO J, 22, 1969-1980.
PDB codes: 1nly 1nlz 1opx
14561776 S.Y.Lee, A.De La Torre, D.Yan, S.Kustu, B.T.Nixon, and D.E.Wemmer (2003).
Regulation of the transcriptional activator NtrC1: structural studies of the regulatory and AAA+ ATPase domains.
  Genes Dev, 17, 2552-2563.
PDB codes: 1ny5 1ny6
14506706 W.Gartner, J.Rossbacher, B.Zierhut, T.Daneva, W.Base, M.Weissel, W.Waldhäusl, M.S.Pasternack, and L.Wagner (2003).
The ATP-dependent helicase RUVBL1/TIP49a associates with tubulin during mitosis.
  Cell Motil Cytoskeleton, 56, 79-93.  
12357035 A.Hansson, R.D.Willows, T.H.Roberts, and M.Hansson (2002).
Three semidominant barley mutants with single amino acid substitutions in the smallest magnesium chelatase subunit form defective AAA+ hexamers.
  Proc Natl Acad Sci U S A, 99, 13944-13949.  
12112691 A.Teplyakov, G.Obmolova, M.Tordova, N.Thanki, N.Bonander, E.Eisenstein, A.J.Howard, and G.L.Gilliland (2002).
Crystal structure of the YjeE protein from Haemophilus influenzae: a putative Atpase involved in cell wall synthesis.
  Proteins, 48, 220-226.
PDB codes: 1fl9 1htw
12006565 A.Zakalskiy, G.Högenauer, T.Ishikawa, E.Wehrschütz-Sigl, F.Wendler, D.Teis, G.Zisser, A.C.Steven, and H.Bergler (2002).
Structural and enzymatic properties of the AAA protein Drg1p from Saccharomyces cerevisiae. Decoupling of intracellular function from ATPase activity and hexamerization.
  J Biol Chem, 277, 26788-26795.  
11782421 D.A.Hattendorf, and S.L.Lindquist (2002).
Cooperative kinetics of both Hsp104 ATPase domains and interdomain communication revealed by AAA sensor-1 mutants.
  EMBO J, 21, 12-21.  
11867765 D.A.Hattendorf, and S.L.Lindquist (2002).
Analysis of the AAA sensor-2 motif in the C-terminal ATPase domain of Hsp104 with a site-specific fluorescent probe of nucleotide binding.
  Proc Natl Acad Sci U S A, 99, 2732-2737.  
11959500 D.Jeruzalmi, M.O'Donnell, and J.Kuriyan (2002).
Clamp loaders and sliding clamps.
  Curr Opin Struct Biol, 12, 217-224.  
12205096 F.Guo, M.R.Maurizi, L.Esser, and D.Xia (2002).
Crystal structure of ClpA, an Hsp100 chaperone and regulator of ClpAP protease.
  J Biol Chem, 277, 46743-46752.
PDB codes: 1k6k 1ksf
11748238 F.X.Gomis-Rüth, G.Moncalían, la Cruz, and M.Coll (2002).
Conjugative plasmid protein TrwB, an integral membrane type IV secretion system coupling protein. Detailed structural features and mapping of the active site cleft.
  J Biol Chem, 277, 7556-7566.  
12377127 H.Niwa, D.Tsuchiya, H.Makyio, M.Yoshida, and K.Morikawa (2002).
Hexameric ring structure of the ATPase domain of the membrane-integrated metalloprotease FtsH from Thermus thermophilus HB8.
  Structure, 10, 1415-1423.
PDB codes: 1ixz 1iy0 1iy1 1iy2
12434150 I.Rouiller, B.DeLaBarre, A.P.May, W.I.Weis, A.T.Brunger, R.A.Milligan, and E.M.Wilson-Kubalek (2002).
Conformational changes of the multifunction p97 AAA ATPase during its ATPase cycle.
  Nat Struct Biol, 9, 950-957.  
12037306 J.Li, and B.Sha (2002).
Cloning, expression, purification and preliminary X-ray crystallographic studies of Escherichia coli Hsp100 nucleotide-binding domain 2 (NBD2).
  Acta Crystallogr D Biol Crystallogr, 58, 1030-1031.  
12234917 J.P.Erzberger, M.M.Pirruccello, and J.M.Berger (2002).
The structure of bacterial DnaA: implications for general mechanisms underlying DNA replication initiation.
  EMBO J, 21, 4763-4773.
PDB code: 1l8q
11909948 K.Miki, and E.M.Eddy (2002).
Tumor necrosis factor receptor 1 is an ATPase regulated by silencer of death domain.
  Mol Cell Biol, 22, 2536-2543.  
12415300 M.J.Davey, D.Jeruzalmi, J.Kuriyan, and M.O'Donnell (2002).
Motors and switches: AAA+ machines within the replisome.
  Nat Rev Mol Cell Biol, 3, 826-835.  
11923293 M.P.Dixon, R.N.Pau, G.J.Howlett, D.E.Dunstan, W.H.Sawyer, and B.E.Davidson (2002).
The central domain of Escherichia coli TyrR is responsible for hexamerization associated with tyrosine-mediated repression of gene expression.
  J Biol Chem, 277, 23186-23192.  
11827963 N.Takahashi, S.Tsutsumi, T.Tsuchiya, B.Stillman, and T.Mizushima (2002).
Functions of sensor 1 and sensor 2 regions of Saccharomyces cerevisiae Cdc6p in vivo and in vitro.
  J Biol Chem, 277, 16033-16040.  
12351638 P.Beinker, S.Schlee, Y.Groemping, R.Seidel, and J.Reinstein (2002).
The N terminus of ClpB from Thermus thermophilus is not essential for the chaperone activity.
  J Biol Chem, 277, 47160-47166.  
12209147 P.Chène (2002).
ATPases as drug targets: learning from their structure.
  Nat Rev Drug Discov, 1, 665-673.  
11959502 X.Zhang, F.Beuron, and P.S.Freemont (2002).
Machinery of protein folding and unfolding.
  Curr Opin Struct Biol, 12, 231-238.  
11340056 A.T.Brunger (2001).
Structure of proteins involved in synaptic vesicle fusion in neurons.
  Annu Rev Biophys Biomol Struct, 30, 157-171.  
11297924 A.T.Brunger (2001).
Structural insights into the molecular mechanism of calcium-dependent vesicle-membrane fusion.
  Curr Opin Struct Biol, 11, 163-173.  
11148030 C.B.Bauer, M.V.Fonseca, H.M.Holden, J.B.Thoden, T.B.Thompson, J.C.Escalante-Semerena, and I.Rayment (2001).
Three-dimensional structure of ATP:corrinoid adenosyltransferase from Salmonella typhimurium in its free state, complexed with MgATP, or complexed with hydroxycobalamin and MgATP.
  Biochemistry, 40, 361-374.
PDB codes: 1g5r 1g5t 1g64
11525729 D.Jeruzalmi, M.O'Donnell, and J.Kuriyan (2001).
Crystal structure of the processivity clamp loader gamma (gamma) complex of E. coli DNA polymerase III.
  Cell, 106, 429-441.
PDB code: 1jr3
11525728 D.Jeruzalmi, O.Yurieva, Y.Zhao, M.Young, J.Stewart, M.Hingorani, M.O'Donnell, and J.Kuriyan (2001).
Mechanism of processivity clamp opening by the delta subunit wrench of the clamp loader complex of E. coli DNA polymerase III.
  Cell, 106, 417-428.
PDB codes: 1jqj 1jql
11250194 G.Mocz, and I.R.Gibbons (2001).
Model for the motor component of dynein heavy chain based on homology to the AAA family of oligomeric ATPases.
  Structure, 9, 93.
PDB code: 1hn5
11296216 I.Hayashi, T.Oyama, and K.Morikawa (2001).
Structural and functional studies of MinD ATPase: implications for the molecular recognition of the bacterial cell division apparatus.
  EMBO J, 20, 1819-1828.
PDB codes: 1g3q 1g3r
11593041 J.T.Littleton, R.J.Barnard, S.A.Titus, J.Slind, E.R.Chapman, and B.Ganetzky (2001).
SNARE-complex disassembly by NSF follows synaptic-vesicle fusion.
  Proc Natl Acad Sci U S A, 98, 12233-12238.  
11709174 J.Wang, J.J.Song, I.S.Seong, M.C.Franklin, S.Kamtekar, S.H.Eom, and C.H.Chung (2001).
Nucleotide-dependent conformational changes in a protease-associated ATPase HsIU.
  Structure, 9, 1107-1116.
PDB codes: 1hqy 1ht1 1ht2
11251810 K.Karata, C.S.Verma, A.J.Wilkinson, and T.Ogura (2001).
Probing the mechanism of ATP hydrolysis and substrate translocation in the AAA protease FtsH by modelling and mutagenesis.
  Mol Microbiol, 39, 890-903.  
11148024 K.Satoh, H.Sasajima, K.I.Nyoumura, H.Yokosawa, and H.Sawada (2001).
Assembly of the 26S proteasome is regulated by phosphorylation of the p45/Rpt6 ATPase subunit.
  Biochemistry, 40, 314-319.  
11171970 K.Yamada, N.Kunishima, K.Mayanagi, T.Ohnishi, T.Nishino, H.Iwasaki, H.Shinagawa, and K.Morikawa (2001).
Crystal structure of the Holliday junction migration motor protein RuvB from Thermus thermophilus HB8.
  Proc Natl Acad Sci U S A, 98, 1442-1447.
PDB code: 1hqc
11389191 M.Mohtashami, B.A.Stewart, G.L.Boulianne, and W.S.Trimble (2001).
Analysis of the mutant Drosophila N-ethylmaleimide sensitive fusion-1 protein in comatose reveals molecular correlates of the behavioural paralysis.
  J Neurochem, 77, 1407-1417.  
11713188 M.R.Maurizi, and C.C.Li (2001).
AAA proteins: in search of a common molecular basis. International Meeting on Cellular Functions of AAA Proteins.
  EMBO Rep, 2, 980-985.  
11309120 M.Su'etsugu, H.Kawakami, K.Kurokawa, T.Kubota, M.Takata, and T.Katayama (2001).
DNA replication-coupled inactivation of DnaA protein in vitro: a role for DnaA arginine-334 of the AAA+ Box VIII motif in ATP hydrolysis.
  Mol Microbiol, 40, 376-386.  
11163235 S.Dalal, and P.I.Hanson (2001).
Membrane traffic: what drives the AAA motor?
  Cell, 104, 5-8.  
11590019 S.J.Russell, F.Gonzalez, L.Joshua-Tor, and S.A.Johnston (2001).
Selective chemical inactivation of AAA proteins reveals distinct functions of proteasomal ATPases.
  Chem Biol, 8, 941-950.  
11473577 T.Ogura, and A.J.Wilkinson (2001).
AAA+ superfamily ATPases: common structure--diverse function.
  Genes Cells, 6, 575-597.  
10851178 A.T.Brunger (2000).
Structural insights into the molecular mechanism of Ca(2+)-dependent exocytosis.
  Curr Opin Neurobiol, 10, 293-302.  
  10790395 B.Dellinger, R.Felling, and R.W.Ordway (2000).
Genetic modifiers of the Drosophila NSF mutant, comatose, include a temperature-sensitive paralytic allele of the calcium channel alpha1-subunit gene, cacophony.
  Genetics, 155, 203-211.  
10871397 C.Venclovas, and M.P.Thelen (2000).
Structure-based predictions of Rad1, Rad9, Hus1 and Rad17 participation in sliding clamp and clamp-loading complexes.
  Nucleic Acids Res, 28, 2481-2493.  
10838059 E.Muneyuki, H.Noji, T.Amano, T.Masaike, and M.Yoshida (2000).
F(0)F(1)-ATP synthase: general structural features of 'ATP-engine' and a problem on free energy transduction.
  Biochim Biophys Acta, 1458, 467-481.  
  10749934 G.J.Steel, C.Harley, A.Boyd, and A.Morgan (2000).
A screen for dominant negative mutants of SEC18 reveals a role for the AAA protein consensus sequence in ATP hydrolysis.
  Mol Biol Cell, 11, 1345-1356.  
10844644 H.Iwasaki, Y.W.Han, T.Okamoto, T.Ohnishi, M.Yoshikawa, K.Yamada, H.Toh, H.Daiyasu, T.Ogura, and H.Shinagawa (2000).
Mutational analysis of the functional motifs of RuvB, an AAA+ class helicase and motor protein for holliday junction branch migration.
  Mol Microbiol, 36, 528-538.  
11163220 I.Rouiller, V.M.Butel, M.Latterich, R.A.Milligan, and E.M.Wilson-Kubalek (2000).
A major conformational change in p97 AAA ATPase upon ATP binding.
  Mol Cell, 6, 1485-1490.  
11208133 J.A.Ybe, D.E.Wakeham, F.M.Brodsky, and P.K.Hwang (2000).
Molecular structures of proteins involved in vesicle fusion.
  Traffic, 1, 474-479.  
11163224 J.Ortega, S.K.Singh, T.Ishikawa, M.R.Maurizi, and A.C.Steven (2000).
Visualization of substrate binding and translocation by the ATP-dependent protease, ClpXP.
  Mol Cell, 6, 1515-1521.  
11114503 K.M.Misura, A.P.May, and W.I.Weis (2000).
Protein-protein interactions in intracellular membrane fusion.
  Curr Opin Struct Biol, 10, 662-671.  
10737940 L.Jaroszewski, L.Rychlewski, J.C.Reed, and A.Godzik (2000).
ATP-activated oligomerization as a mechanism for apoptosis regulation: fold and mechanism prediction for CED-4.
  Proteins, 39, 197-203.  
11106733 M.C.Sousa, C.B.Trame, H.Tsuruta, S.M.Wilbanks, V.S.Reddy, and D.B.McKay (2000).
Crystal and solution structures of an HslUV protease-chaperone complex.
  Cell, 103, 633-643.
PDB codes: 1g3i 1g3k
10893253 R.D.Vale (2000).
AAA proteins. Lords of the ring.
  J Cell Biol, 150, F13-F19.  
10966477 T.J.Kelly, and G.W.Brown (2000).
Regulation of chromosome replication.
  Annu Rev Biochem, 69, 829-880.  
10782097 T.Langer (2000).
AAA proteases: cellular machines for degrading membrane proteins.
  Trends Biochem Sci, 25, 247-251.  
11163219 X.Zhang, A.Shaw, P.A.Bates, R.H.Newman, B.Gowen, E.Orlova, M.A.Gorman, H.Kondo, P.Dokurno, J.Lally, G.Leonard, H.Meyer, M.van Heel, and P.S.Freemont (2000).
Structure of the AAA ATPase p97.
  Mol Cell, 6, 1473-1484.
PDB code: 1e32
10809689 Y.Shotland, A.Shifrin, T.Ziv, D.Teff, S.Koby, O.Kobiler, and A.B.Oppenheim (2000).
Proteolysis of bacteriophage lambda CII by Escherichia coli FtsH (HflB).
  J Bacteriol, 182, 3111-3116.  
10368290 A.F.Neuwald (1999).
The hexamerization domain of N-ethylmaleimide-sensitive factor: structural clues to chaperone function.
  Structure, 7, R19-R23.  
10580154 A.K.Pullikuth, and S.S.Gill (1999).
Identification of a Manduca sexta NSF ortholog, a member of the AAA family of ATPases.
  Gene, 240, 343-354.  
10359771 C.K.Smith, T.A.Baker, and R.T.Sauer (1999).
Lon and Clp family proteases and chaperones share homologous substrate-recognition domains.
  Proc Natl Acad Sci U S A, 96, 6678-6682.  
10404598 D.Fass, C.E.Bogden, and J.M.Berger (1999).
Crystal structure of the N-terminal domain of the DnaB hexameric helicase.
  Structure, 7, 691-698.
PDB code: 1b79
10872471 D.Voges, P.Zwickl, and W.Baumeister (1999).
The 26S proteasome: a molecular machine designed for controlled proteolysis.
  Annu Rev Biochem, 68, 1015-1068.  
10531065 J.J.Hartman, and R.D.Vale (1999).
Microtubule disassembly by ATP-dependent oligomerization of the AAA enzyme katanin.
  Science, 286, 782-785.  
10473576 K.Karata, T.Inagawa, A.J.Wilkinson, T.Tatsuta, and T.Ogura (1999).
Dissecting the role of a conserved motif (the second region of homology) in the AAA family of ATPases. Site-directed mutagenesis of the ATP-dependent protease FtsH.
  J Biol Chem, 274, 26225-26232.  
10340698 L.M.Quarmby, and T.A.Lohret (1999).
Microtubule severing.
  Cell Motil Cytoskeleton, 43, 1-9.  
10445030 L.M.Rice, and A.T.Brunger (1999).
Crystal structure of the vesicular transport protein Sec17: implications for SNAP function in SNARE complex disassembly.
  Mol Cell, 4, 85-95.
PDB code: 1qqe
10531028 M.Coles, T.Diercks, J.Liermann, A.Gröger, B.Rockel, W.Baumeister, K.K.Koretke, A.Lupas, J.Peters, and H.Kessler (1999).
The solution structure of VAT-N reveals a 'missing link' in the evolution of complex enzymes from a simple betaalphabetabeta element.
  Curr Biol, 9, 1158-1168.
PDB codes: 1cz4 1cz5
10508673 M.Schmidt, A.N.Lupas, and D.Finley (1999).
Structure and mechanism of ATP-dependent proteases.
  Curr Opin Chem Biol, 3, 584-591.  
10473546 P.Zwickl, D.Ng, K.M.Woo, H.P.Klenk, and A.L.Goldberg (1999).
An archaebacterial ATPase, homologous to ATPases in the eukaryotic 26 S proteasome, activates protein breakdown by 20 S proteasomes.
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10582236 P.Zwickl, D.Voges, and W.Baumeister (1999).
The proteasome: a macromolecular assembly designed for controlled proteolysis.
  Philos Trans R Soc Lond B Biol Sci, 354, 1501-1511.  
10445031 R.C.Yu, R.Jahn, and A.T.Brunger (1999).
NSF N-terminal domain crystal structure: models of NSF function.
  Mol Cell, 4, 97.
PDB code: 1qcs
10872468 R.Jahn, and T.C.Südhof (1999).
Membrane fusion and exocytosis.
  Annu Rev Biochem, 68, 863-911.  
10611286 S.M.Babor, and D.Fass (1999).
Crystal structure of the Sec18p N-terminal domain.
  Proc Natl Acad Sci U S A, 96, 14759-14764.
PDB code: 1cr5
9875840 J.J.Skehel, and D.C.Wiley (1998).
Coiled coils in both intracellular vesicle and viral membrane fusion.
  Cell, 95, 871-874.  
9783736 J.Rizo, and T.C.Südhof (1998).
Mechanics of membrane fusion.
  Nat Struct Biol, 5, 839-842.  
9822570 R.Jahn (1998).
Synaptic transmission: two players team up for a new tune.
  Curr Biol, 8, R856-R858.  
9844627 T.M.Hohl, F.Parlati, C.Wimmer, J.E.Rothman, T.H.Söllner, and H.Engelhardt (1998).
Arrangement of subunits in 20 S particles consisting of NSF, SNAPs, and SNARE complexes.
  Mol Cell, 2, 539-548.  
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