FB2024_03 , released June 25, 2024
Gene: Dmel\car
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General Information
Symbol
Dmel\car
Species
D. melanogaster
Name
carnation
Annotation Symbol
CG12230
Feature Type
FlyBase ID
FBgn0000257
Gene Model Status
Stock Availability
Gene Summary
carnation (car) encodes a subunit of the conserved HOPS complex, which is required for pigment granule formation and membrane fusion events, such as fusions between endosomes or autophagosomes with lysosomes. car loss of function mutant flies exhibit altered eye color. [Date last reviewed: 2018-09-06] (FlyBase Gene Snapshot)
Also Known As

Vps33, Vps33a, l(1)G0447

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-63
RefSeq locus
NC_004354 REGION:19566019..19569401
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (27 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
contributes_to syntaxin binding
inferred from physical interaction with FLYBASE:Syx17; FB:FBgn0035540
inferred from physical interaction with FLYBASE:Syx16; FB:FBgn0031106
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence or structural similarity
Biological Process (19 terms)
Terms Based on Experimental Evidence (14 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:p; FB:FBgn0086679
involved_in crinophagy
inferred from mutant phenotype
inferred from mutant phenotype
involved_in endocytosis
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
traceable author statement
inferred from sequence or structural similarity with SGD:S000004388
inferred from biological aspect of ancestor with PANTHER:PTN000187655
traceable author statement
inferred from sequence or structural similarity with SGD:S000004388
inferred from electronic annotation with InterPro:IPR001619
inferred from sequence or structural similarity
inferred from biological aspect of ancestor with PANTHER:PTN000187655
Cellular Component (5 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from direct assay
part_of HOPS complex
inferred from mutant phenotype
inferred from physical interaction with UniProtKB:Q24314, UniProtKB:Q5KU39, UniProtKB:Q9VHG1
inferred from direct assay
located_in lysosome
inferred from direct assay
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000187807
inferred from sequence or structural similarity with SGD:S000004388
located_in cytosol
traceable author statement
part_of HOPS complex
inferred from sequence or structural similarity with SGD:S000004388
is_active_in lysosome
inferred from biological aspect of ancestor with PANTHER:PTN000913042
located_in membrane
traceable author statement
Protein Family (UniProt)
Belongs to the STXBP/unc-18/SEC1 family. (Q9Y1I2)
Summaries
Gene Snapshot
carnation (car) encodes a subunit of the conserved HOPS complex, which is required for pigment granule formation and membrane fusion events, such as fusions between endosomes or autophagosomes with lysosomes. car loss of function mutant flies exhibit altered eye color. [Date last reviewed: 2018-09-06]
Gene Group (FlyBase)
SM TRAFFICKING GENES -
Members of the Sec1/Munc18 (SM) gene family interact with soluble N-ethylmaleimide-sensitive factor attachment protein receptors (SNAREs) to regulate SNARE complex assembly and vesicle membrane fusion. (Adapted from PMID:20471239).
VPS-C COMPLEXES: HOPS AND CORVET -
The heterohexameric VPS-C complexes, HOPS and CORVET have been extensively characterised in yeast. They interact with SNAREs and Rabs to promote vesicle tethering. The CORVET complex is involved in early endosome fusion, interacting with Rab5-GTP. The HOPS complex interacts with Rab7-GTP on late endosomes (multivesicular bodies) to promote fusion with the lysosomal compartment (and vacuoles) and also autophagosome-lysosome fusion. The VPS-C complexes share common subunits and CORVET to HOPS conversion intermediates (i-CORVET and i-HOPS) have been proposed to accompany early to late endosome maturation. (Adapted from PMID:19575650 and FBrf0223135).
Protein Function (UniProtKB)
Core component of the class C core vacuole/endosome tethering (CORVET) and the homotypic fusion and vacuole protein sorting (HOPS) tethering complexes involved in endo-lysosomal vesicle trafficking and lysosome biogenesis (PubMed:10549280, PubMed:22160599, PubMed:24554766, PubMed:27253064). The CORVET complex facilitates docking and fusion of endosomal vesicles during endosome maturation, acts upstream of HOPS, but is not involved in autophagic flux (PubMed:10549280, PubMed:27253064, PubMed:31194677). The CORVET complex may cooperate with the early endosomal tether Rbsn-5 to mediate endosomal fusion (PubMed:27253064). The HOPS complex facilitates docking and fusion of lysosomes with late endosomes and several other types of vesicles (PubMed:24554766, PubMed:31194677). The HOPS complex is also involved in autophagy and crinophagy (the elimination of unused secretory granules through their fusion with lysosomes) (PubMed:24554766, PubMed:31194677). The HOPS complex probably instigates autophagosome-lysosome fusion by binding autophagosome associated Syx17/syntaxin 17 and promoting assembly of the trans-SNARE complex (PubMed:24554766). Independent of Syx17/syntaxin 17 HOPS is involved in biosynthetic transport to lysosomes and lysosome-related organelles such as eye-pigment granules (PubMed:10549280, PubMed:24554766, PubMed:31194677). Required for endocytic degradation of boss/bride of sevenless and N/Notch in developing ommatidia (PubMed:24554766).
(UniProt, Q9Y1I2)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
car: carnation
Eye color dark ruby. Body shape and proportions seem rounded. With st, eye color is yellow-brown; with bw, brownish yellow to brown (Mainx, 1938, Z. Indukt. Abstamm. Vererbungsl. 75: 256-76). Malpighian tubes pale yellow in mature larva (Beadle, 1937, Genetics 22: 587-611) but hard to distinguish from wild type before third instar. Eye color autonomous in transplant into wild-type host (Beadle and Ephrussi, 1936, Genetics 21: 230). car dor is pupal lethal and shows reduced recovery in gynandromorphs with male parts car dor (Nash, 1971, DIS 47: 73). car lt also lethal, dies as larva when mother car;lt/+ or car/+;lt and as pupa when mother car/+;lt/+ (Nickla, 1977, Nature 268: 638-39); lethal focus domineering; fate maps to ventral nervous system (Nickla, Lilly, and McCarthy, 1980, Experientia 36: 402-05). Brain histology abnormal [McCarthy and Nickla, 1980, Experientia 36: 1361-63 (fig.)].
Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\car 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 Q9Y1I2)

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 5.40

Gene model reviewed during 5.44

Stop-codon suppression (UGA) postulated; FBrf0216884.

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)
FBtr0074728
3097
617
FBtr0074727
3173
617
FBtr0331158
3173
639
Additional Transcript Data and Comments
Reported size (kB)

3.022 (longest cDNA)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0074497
68.8
617
6.74
FBpp0074496
68.8
617
6.74
FBpp0303585
71.2
639
7.13
Polypeptides with Identical Sequences

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

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

Component of the class C core vacuole/endosome tethering (CORVET) complex composed of at least Vps8, dor/Vps18, car/Vps33A and Vps16A; unlike in other species, Vps11 is not part of the Drosophila complex (PubMed:10549280, PubMed:27253064). Due to the reduced number of components the Drosophila CORVET complex is often referred to as the miniCORVET complex (PubMed:27253064). Interacts with ema (PubMed:20194640). Component of the homotypic fusion and vacuole protein sorting (HOPS) complex, composed of Vps16A, car/Vps33A, dor/Vps18, Vps39, Vps11 and lt/Vps41 (PubMed:31194677). The tethering complex core made up of Vps16A, car/Vps33A and dor/Vps18 and shared by both HOPS and CORVET, preferentially associates with CORVET specific Vps8 over HOPS specific lt/Vps41 (PubMed:31194677). Interacts with Syx17 (via SNARE domain); the interaction requires Vps16A, may involve additional components of the HOPS complex and may promote assembly of the Syx17-Snap29-Vamp7 trans-SNARE complex (PubMed:24554766).

(UniProt, Q9Y1I2)
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\car using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
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).

1.51

Transcript Expression
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
part_of HOPS complex
inferred from mutant phenotype
inferred from physical interaction with UniProtKB:Q24314, UniProtKB:Q5KU39, UniProtKB:Q9VHG1
inferred from direct assay
located_in lysosome
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\car 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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
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
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 10 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of car
Transgenic constructs containing regulatory region of car
Aberrations (Deficiencies and Duplications) ( 37 )
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
Sterility
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) (7)
14 of 14
Yes
Yes
6 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1  
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (7)
13 of 14
Yes
Yes
6 of 14
No
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
Mus musculus (laboratory mouse) (7)
13 of 14
Yes
Yes
6 of 14
No
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
Xenopus tropicalis (Western clawed frog) (9)
7 of 13
Yes
Yes
3 of 13
No
No
3 of 13
No
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
Danio rerio (Zebrafish) (8)
13 of 14
Yes
Yes
6 of 14
No
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
2 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (6)
10 of 14
Yes
Yes
8 of 14
No
Yes
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (6)
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (9)
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
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (4)
10 of 13
Yes
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (4)
12 of 12
Yes
Yes
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:car. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (4)
5 of 13
4 of 13
4 of 13
3 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
esyN Network Diagram
Show/hide secondary interactors 
(data from AllianceMine provided by esyN)
esyN Network Key:
Suppression
Enhancement
Other 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)
Component of the class C core vacuole/endosome tethering (CORVET) complex composed of at least Vps8, dor/Vps18, car/Vps33A and Vps16A; unlike in other species, Vps11 is not part of the Drosophila complex (PubMed:10549280, PubMed:27253064). Due to the reduced number of components the Drosophila CORVET complex is often referred to as the miniCORVET complex (PubMed:27253064). Interacts with ema (PubMed:20194640). Component of the homotypic fusion and vacuole protein sorting (HOPS) complex, composed of Vps16A, car/Vps33A, dor/Vps18, Vps39, Vps11 and lt/Vps41 (PubMed:31194677). The tethering complex core made up of Vps16A, car/Vps33A and dor/Vps18 and shared by both HOPS and CORVET, preferentially associates with CORVET specific Vps8 over HOPS specific lt/Vps41 (PubMed:31194677). Interacts with Syx17 (via SNARE domain); the interaction requires Vps16A, may involve additional components of the HOPS complex and may promote assembly of the Syx17-Snap29-Vamp7 trans-SNARE complex (PubMed:24554766).
(UniProt, Q9Y1I2 )
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Signaling Pathways (FlyBase)
Metabolic Pathways
External Data
Linkouts
KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
Genomic Location and Detailed Mapping Data
Chromosome (arm)
X
Recombination map
1-63
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
18D1-18D1
Limits computationally determined from genome sequence between P{EP}EP1526&P{EP}PfrxEP1150 and P{EP}Tao-1EP1455
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
18D1-18D4
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Location

1-62.5

Notes
Stocks and Reagents
Stocks (51)
Genomic Clones (20)
cDNA Clones (72)
 

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
    DRSC - Results frm RNAi screens
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
        Other Comments

        car is required for endocytic trafficking to lysosomes. car is required for the biogenesis of pigment granules.

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

        dor is part of a protein complex which includes car.

        Lesions in car reduce eye and ocelli pigmentation and cause excretion of pigment from the tubules.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: car Vps33

        Source for merge of: car l(1)G0447

        Additional comments

        "l(1)G0041" may affect "car".

        Nomenclature History
        Source for database identify of
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (16)
        Reported As
        Symbol Synonym
        Vps33/carnation
        Secondary FlyBase IDs
        • FBgn0027357
        • FBgn0040160
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 51 )
        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.
        BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
        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
        FlyMine - An integrated database for Drosophila genomics
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
        DroID - A comprehensive database of gene and protein interactions.
        DRSC - Results frm RNAi screens
        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        Flygut - An atlas of the Drosophila adult midgut
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (151)