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General Information
Symbol
Dmel\TER94
Species
D. melanogaster
Name
Annotation Symbol
CG2331
Feature Type
FlyBase ID
FBgn0286784
Gene Model Status
Stock Availability
Enzyme Name (EC)
Adenosinetriphosphatase (3.6.1.3)
Gene Snapshot
NA (TER94) encodes a protein that operates as a hexamere, functioning as a ubiquitin-dependent molecular chaperone. The hexamere associates with different co-factors to process nuclear and cytoplasmic substrates marked by ubiquitin. [Date last reviewed: 2019-09-26]
Also Known As
VCP, p97, dVCP, l(2)03775, dmTERA
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:9,989,159..9,993,559 [+]
Recombination map
2-61
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the AAA ATPase family. (Q7KN62)
Molecular Function (GO)
[Detailed GO annotations]
Catalytic Activity (EC)
Experimental Evidence
-
Predictions / Assertions
ATP + H(2)O = ADP + phosphate (3.6.1.3)
Summaries
Pathway (FlyBase)
Negative Regulators of Wnt-TCF Signaling Pathway -
Negative regulators of canonical Wnt signaling down-regulate the pathway, resulting in the attenuation of transcriptional regulation mediated by β-catenin (arm).
Protein Function (UniProtKB)
Necessary for the fragmentation of Golgi stacks during mitosis and for their reassembly after mitosis. Involved in the formation of the transitional endoplasmic reticulum (tER). The transfer of membranes from the endoplasmic reticulum to the Golgi apparatus occurs via 50-70 nm transition vesicles which derive from part-rough, part-smooth transitional elements of the endoplasmic reticulum (tER). Vesicle budding from the tER is an ATP-dependent process. Involved in the ubiquitin-proteasome system. Important for oskar mRNA localization and/or anchoring during oogenesis. Involved in germ cell formation.
(UniProt, Q7KN62)
Summary (Interactive Fly)
AAA family ATPase, CDC48 subfamily - modulation of proteolytic degradation - Hedgehog pathway -Wingless pathway - dendritic pruning -motor neuron degeneration - ER stress response - maintenance of paternal chromosome integrity in the Drosophila zygote
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
4

Please see the GBrowse view of Dmel\TER94 or the JBrowse view of Dmel\TER94 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
Comments on Gene Model
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.46
Gene model reviewed during 5.52
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0088391
3016
801
FBtr0112905
3058
826
FBtr0112906
3125
759
FBtr0343852
3055
825
Additional Transcript Data and Comments
Reported size (kB)
3.2 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0087479
88.9
801
4.98
FBpp0111818
92.1
826
5.00
FBpp0111819
84.2
759
4.99
FBpp0310395
91.9
825
5.00
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
799 (aa); 94 (kD predicted)
94 (kD observed); 89 (kD predicted)
801 (aa); 94 (kD observed)
Comments
TER94 protein from ovarian extracts sediments at close to the expected size for a homohexameric complex. Such complexes are also observed for vertebrate TERs and yeast CDC48.
External Data
Subunit Structure (UniProtKB)
Homohexamer (PubMed:10564274). Interacts with tud, vas, papi and AGO3 (PubMed:18590813, PubMed:21447556). Interacts with Npl4 (PubMed:26471729).
(UniProt, Q7KN62)
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\TER94 using the Feature Mapper tool.

External Data
Crossreferences
Linkouts
Gene Ontology (40 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from physical interaction with UniProtKB:Q9VBP9
(assigned by UniProt )
inferred from physical interaction with FLYBASE:papi; FB:FBgn0031401
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR003959, InterPro:IPR003960
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN000554398
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
Biological Process (26 terms)
Terms Based on Experimental Evidence (20 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:vas; FB:FBgn0283442
inferred from genetic interaction with FLYBASE:tud; FB:FBgn0003891
inferred from mutant phenotype
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from sequence or structural similarity with SGD:S000002284
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
Cellular Component (10 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
inferred from high throughput direct assay
inferred from high throughput direct assay
inferred from direct assay
inferred from high throughput direct assay
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000554506
(assigned by GO_Central )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism | ubiquitous

Comment: maternally deposited

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
TER94 transcripts are strongly expressed in nurse cells starting in stage 9-10 of oogenesis. Strong expression is also observed in developing ommatidia in the eye imaginal disc.
TER94 transcripts are present in germarium region 1 and are present in all germ line cells and follicle cells.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
TER94 protein is present predominantly in germarium region 2b and in stage 1 egg chambers.
TER94 protein is observed in ovarian germ cells and somatic cells.
TER94 protein is largely cytoplasmic and is enriched in fusomes. A low level of protein is observed in germ cell nuclei. Strong nuclear staining was observed in nonovarian somatic cells in larvae and adults.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
inferred from high throughput direct assay
inferred from high throughput direct assay
inferred from direct assay
inferred from high throughput direct assay
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\TER94 in GBrowse 2
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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 16 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 34 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of TER94
Transgenic constructs containing regulatory region of TER94
Deletions and Duplications ( 19 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
endoplasmic reticulum & egg chamber
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (6)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
 
 
1 of 15
No
Yes
1 of 15
No
Yes
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
1 of 15
No
Yes
1 of 15
No
No
Rattus norvegicus (Norway rat) (5)
13 of 13
Yes
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (3)
10 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (3)
14 of 15
Yes
Yes
12 of 15
No
Yes
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
13 of 15
Yes
Yes
13 of 15
Yes
Yes
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (4)
9 of 9
Yes
Yes
9 of 9
Yes
Yes
9 of 9
Yes
Yes
1 of 9
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (2)
15 of 15
Yes
Yes
1 of 15
No
No
Schizosaccharomyces pombe (Fission yeast) (1)
12 of 12
Yes
Yes
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091902Q2 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091501PU )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W01KH )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X01C9 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G024K )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (5)
3 of 10
2 of 10
2 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 6 )
Potential Models Based on Orthology ( 3 )
Modifiers Based on Experimental Evidence ( 11 )
Allele
Disease
Interaction
References
Comments on Models/Modifiers Based on Experimental Evidence ( 4 )
 
Eye degeneration used to model disease. Point mutation in TER94A229E.Scer\UAS.R models that associated with the most severe form of Inclusion Body Myopathy associated with Paget’s Disease of Bone
Eye degeneration used to model disease. Point mutation in TER94R152H.Scer\UAS.R models that most commonly found in Inclusion Body Myopathy associated with Paget’s Disease of Bone
and Frontotemporal Dementia.
Contrary to expection, TER9426-8 does not suppress polyQ-induced neurodegeneration by inhibiting polyQ aggregation.
Disease Associations of Human Orthologs (via DIOPT v7.1 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.
Dmel gene
Ortholog showing functional complementation
Supporting References
Interactions
Summary of Physical Interactions
esyN Network Diagram
Show neighbor-neighbor interactions:
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Interactions Browser

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

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 (PubMed:10564274). Interacts with tud, vas, papi and AGO3 (PubMed:18590813, PubMed:21447556). Interacts with Npl4 (PubMed:26471729).
(UniProt, Q7KN62 )
Linkouts
BioGRID - A database of protein and genetic interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
Pathways
Gene Group - Pathway Membership (FlyBase)
Negative Regulators of Wnt-TCF Signaling Pathway -
Negative regulators of canonical Wnt signaling down-regulate the pathway, resulting in the attenuation of transcriptional regulation mediated by β-catenin (arm).
External Data
Linkouts
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2R
Recombination map
2-61
Cytogenetic map
Sequence location
2R:9,989,159..9,993,559 [+]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
46D1-46E2
Limits computationally determined from genome sequence between P{lacW}Adamk13906&P{EP}Pka-R2EP2162 and P{PZ}14-3-3ζ07103; Left limit from non-inclusion within Df(2R)B5 (FBrf0074052) Right limit from inclusion within Df(2R)X1 (FBrf0074052)
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
The l(2)46Db complementation group maps to the genomic interval 2R:9959818..10025288 , release 6 genome (left limit from complementation mapping against Df(2R)01D09Y-M073, right limit from complementation mapping against Df(2R)BSC158).
Complementation data from unspecified deficiency chromosomes.
46D-46D
(determined by in situ hybridisation)
46D1-46D2
(determined by in situ hybridisation)
46C-46D
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (19)
Genomic Clones (17)
 

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

cDNA Clones (201)
 

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 GBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequences
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
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: TER94 l(2)03775
    Source for merge of: TER94 Apple
    Source for merge of: TER94 l(2)46Ch
    Source for merge of: TER94 l(2)46CFf l(2)46CFs
    Source for merge of: Tera TER94
    Source for merge of: TER94 l(2)46Db
    Additional comments
    The l(2)46Db complementation group corresponds to TER94, even though l(2)46Db[2] shows a complex complementation pattern with other TER94 alleles.
    Other Comments
    TER94-dependent th degradation is required for proper neuronal remodelling and apoptosis.
    TER94 inhibition interferes with class IV neuron dendrite pruning and class III neuron apoptosis during the pupal stage.
    dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
    TER94 is involved in osk localisation during oogenesis.
    Identification: approximately 3.3kb transcript identified during molecular analysis of the eve genomic region.
    Identification: Physical interaction with bam protein in a yeast two hybrid assay.
    TER94 has been cloned using an antibody to Canp, a Ca2+-activated protease (FBrf0055463), but TER94 protein expressed in E.coli does not show Ca2+-activated protease activity.
    A Drosophila homolog of CDC48, a protein that has been linked to vesicle fusion in yeast and vertebrate cells, isolated in a protein interaction screen showing interaction with bam.
    Complementation group identified in an EMS and DEB screen to isolate deficiencies that uncover Jra.
    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 56 )
    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/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
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    Synonyms and Secondary IDs (47)
    Reported As
    Symbol Synonym
    BcDNA.GM02885
    BcDNA:GM02885
    Tera
    anon-WO2004063362.65
    anon-WO2004063362.67
    l(2)46CFf
    l(2)46CFs
    l(2)46Ch
    Name Synonyms
    Complementation group I
    TRANSITIONAL ENDOPLASMIC RETICULUM ATPASE TER94
    Transitional Endoplasmic Reticulum 94
    Transitional endoplasmic reticulum 94
    Transitional endoplasmic reticulum ATPase
    Valosin containing protein
    Valosin-Containing Protein
    lethal (2) 46Db
    transitional endoplasmic reticulum ATPase
    valosin-containing protein
    Secondary FlyBase IDs
    • FBgn0014986
    • FBgn0261014
    • FBgn0024923
    • FBgn0010554
    • FBgn0015484
    • FBgn0019709
    • FBgn0019722
    • FBgn0025353
    • FBgn0027618
    • FBgn0064213
    • FBgn0082213
    • FBgn0082214
    • FBgn0024280
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (194)