FB2025_05 , released December 11, 2025
Gene: Dmel\Nsf2
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General Information
Symbol
Dmel\Nsf2
Species
D. melanogaster
Name
N-ethylmaleimide-sensitive factor 2
Annotation Symbol
CG33101
Feature Type
FlyBase ID
FBgn0290174
Gene Model Status
Stock Availability
Gene Summary
N-ethylmaleimide-sensitive factor 2 (Nsf2) encodes a homohexameric AAA ATPase that functions to recycle SNARE complex components subsequent to membrane fusion. In the tracheal system, it is required for tube growth and connectivity within the terminal cell. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

dNSF2, dNSF-2, urd, NEM-sensitive fusion protein 2, lotus

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-54
RefSeq locus
NT_033777 REGION:13833497..13837991
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (3 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Biological Process (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR039812
Cellular Component (0 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (0 terms)
Protein Family (UniProt)
Belongs to the AAA ATPase family. (P54351)
Catalytic Activity (EC/Rhea)
ATP hydrolysis activity
RHEA 13065:
Summaries
Gene Snapshot
N-ethylmaleimide-sensitive factor 2 (Nsf2) encodes a homohexameric AAA ATPase that functions to recycle SNARE complex components subsequent to membrane fusion. In the tracheal system, it is required for tube growth and connectivity within the terminal cell. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
N-ETHYLMALEIMIDE-SENSITIVE FACTORS -
The N-ethylmaleimide-sensitive factor/fusion protein (NSF), is a hexameric ATPase recruited to SNARE-SNAP complexes. ATP hydrolysis by NSF, leads to conformational changes and dissociation of the SNARE complex. (Adapted from FBrf0141270).
Protein Function (UniProtKB)
Required for vesicle-mediated transport. Catalyzes the fusion of transport vesicles within the Golgi cisternae. Is also required for transport from the endoplasmic reticulum to the Golgi stack. Seems to function as a fusion protein required for the delivery of cargo proteins to all compartments of the Golgi stack independent of vesicle origin (By similarity).
(UniProt, P54351)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
urd: urdur (J.A. Kennison)
Isolated as a dominant suppressor of Pc mutations. Also suppresses Pcl and Msc alleles. Recessive larval lethal.
Summary (Interactive Fly)

homohexameric AAA ATPase - developmentally regulated N-ethylmaleimide-sensitive fusion (NSF) gene - mutants are defective in the synaptic growth and maturation that occurs during larval development - functions to recycle SNARE complex components subsequent to membrane fusion - required for tracheal tube growth and connectivity within the terminal cell

Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Nsf2 for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P54351)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.55

Low-frequency RNA-Seq exon junction(s) not annotated.

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0082883
2427
752
FBtr0344766
2823
752
FBtr0344767
3521
752
Additional Transcript Data and Comments
Reported size (kB)

2.4 (northern blot)

3.4, 3.0, 2.8, 2.4 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0082346
83.4
752
6.71
FBpp0311094
83.4
752
6.71
FBpp0311095
83.4
752
6.71
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

752 aa isoforms: Nsf2-PA, Nsf2-PB, Nsf2-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Homohexamer.

(UniProt, P54351)
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Nsf2 using the Feature Mapper tool.

External Data
Crossreferences
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

2.73

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

The highest levels of Nsf2 transcripts are seen before and during cellularization. After gastrulation, expression is seen primarily in regions which give rise to ectodermal and endodermal tissues such as the midgut and the hindgut. Later, expression is also seen in the CNS. In larvae, Nsf2 expression is observed in the ring gland and in a restricted number of cells in the salivary gland. Expression is also observed in unusual patterns in the imaginal discs with the highest levels in the leg, wing and antennal discs.

Nsf2 transcripts are expressed at substantially higher levels in adults and larvae than in embryos.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Nsf2 in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 20 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Nsf2
Transgenic constructs containing regulatory region of Nsf2
Aberrations (Deficiencies and Duplications) ( 3 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (24)
14 of 14
Yes
Yes
8  
2 of 14
No
No
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1  
2 of 14
No
No
2  
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
10  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
2  
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (14)
13 of 14
Yes
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (15)
13 of 14
Yes
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (20)
7 of 13
Yes
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (18)
14 of 14
Yes
Yes
13 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (19)
13 of 14
Yes
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (25)
11 of 12
Yes
Yes
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (48)
11 of 13
Yes
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (16)
13 of 13
Yes
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (9)
12 of 12
Yes
Yes
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Nsf2. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (24)
12 of 13
6 of 13
2 of 13
2 of 13
2 of 13
2 of 13
2 of 13
2 of 13
2 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 1 )
Modifiers Based on Experimental Evidence ( 2 )
Allele
Disease
Interaction
References
Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Functional Complementation Data
Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
Interactions
Summary of Physical Interactions
Summary of Genetic Interactions
Interaction Browsers

Please look at the allele data for full details of the genetic interactions
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
External Data
Subunit Structure (UniProtKB)
Homohexamer.
(UniProt, P54351 )
Linkouts
Pathways
Signaling Pathways (FlyBase)
Metabolic Pathways
FlyBase
External Links
External Data
Linkouts
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-54
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
87F15-87F15
Limits computationally determined from genome sequence between P{PZ}flfl01949 and P{lacW}trxj14A6&P{PZ}trx00347; Left limit from inclusion within Df(3R)red3l (FBrf0102846) Right limit from inclusion within Df(3R)urd (citation unavailable)
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Determined by comparing Celera genomic sequence with sequence from BDGP BAC and P1 clones.
87F12-88A3
87F12-87F15
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (13)
Genomic Clones (17)
 

Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

cDNA Clones (153)
 

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequenced
BDGP DGC clones
Other clones
    Drosophila Genomics Resource Center cDNA clones

    For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

      cDNA Clones, End Sequenced (ESTs)
      RNAi and Array Information
      Linkouts
      Antibody Information
      Laboratory Generated Antibodies
       
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          ChEST reveals this is a target of Mef2.

          dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

          Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days. Shows significant change of expression pattern in circadian mutant background; decreased expression in per01, tim01 and increased expression in ClkJrk background.

          comt and Nsf2 exhibit similar functional properties but have evolved distinct tissue-specific roles. comt is required in the nervous system beginning at the adult stage of development whereas Nsf2 is required in the mesoderm beginning at the first larval instar. The comt and Nsf2 proteins can functionally substitute for one another.

          Evidence of multiple, highly related isoforms of NSF in a single species suggests that in some tissue membranes fusion and/or secretion has a specific role during development.

          Sections of the Scr regulatory region may be important for regulation of Scr by Polycomb- and trithorax-group genes.

          Drosophila has a second N-ethylmaleimide-sensitive fusion protein (NSF) homolog, Nsf2. comt and Nsf2 display overlapping but different temporal expression patterns. Multiple transcripts derived from Nsf2.

          urd is one of the 18 loci identified in a screen for dominant modifiers of Pc and/or Antp phenotypes. Cytological map position stated to be 87F12-15, derived from complementation analysis with deficiency and duplication chromosomes.

          Isolated as a dominant suppressor of Pc mutations. Also suppresses Pcl and Msc alleles.

          Relationship to Other Genes
          Source for database merge of

          Source for merge of: Nsf2 urd lotus

          Additional comments

          Annotation CG9931 split into CG33101 (Nsf2) and CG31495 in release 3 of the genome annotation.

          Nomenclature History
          Source for database identify of

          Source for identity of: Nsf2 CG9931

          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (19)
          Reported As
          Symbol Synonym
          Name Synonyms
          N-ethylmaleimide sensitive factor 2
          N-ethylmaleimide-sensitive factor 2
          urdur
          Secondary FlyBase IDs
          • FBgn0266464
          • FBgn0264653
          • FBgn0013998
          • FBgn0053101
          • FBgn0003960
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 36 )
          Sequence Crossreferences
          NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
          GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
          GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
          RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
          UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
          UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
          UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
          Other crossreferences
          AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
          DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
          EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
          FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          Linkouts
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          References (144)