FB2026_02 , released June 18, 2026
FB2026_02 , released June 18, 2026
Gene: Dmel\cav
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General Information
Symbol
Dmel\cav
Species
D. melanogaster
Name
caravaggio
Annotation Symbol
CG6219
Feature Type
FlyBase ID
FBgn0026257
Gene Model Status
Stock Availability
Gene Summary
caravaggio (cav) is a fast evolving gene that encodes HP1/ORC Associated Protein (HOAP). HOAP is a DNA binding protein strongly enriched at chromosome ends and required for telomere capping. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
Also Known As

HOAP, anon1G5, HP1/ORC-associated protein, p55, anon fe 1G5

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-84
RefSeq locus
NT_033777 REGION:24221103..24223528
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (9 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with UniProtKB:B7Z0L8
inferred from physical interaction with UniProtKB:Q9VR09
inferred from physical interaction with UniProtKB:P05205
Terms Based on Predictions or Assertions (0 terms)
Biological Process (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
involved_in telomere capping
inferred from expression pattern
inferred from mutant phenotype
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Summaries
    Gene Snapshot
    caravaggio (cav) is a fast evolving gene that encodes HP1/ORC Associated Protein (HOAP). HOAP is a DNA binding protein strongly enriched at chromosome ends and required for telomere capping. [Date last reviewed: 2019-03-07]
    Gene Group (FlyBase)
    HIPHOP-HOAP TELOMERE-CAPPING COMPLEX -
    The HipHop-HOAP telomere-capping complex binds to the double stranded portion of telomeric chromatin and recruits Su(var)205 (Heterochromatin Protein 1), a structural component of heterochromatin involved in gene repression, to telomeres. Also recruits the MTV complex to form the terminin telomere-capping complex, which binds to chromosome ends in a sequence-independent manner and prevents telomere fusion. (Adapted from ComplexPortal.)
    MTV COMPLEX -
    The Moi-Tea-Ver (MTV) complex binds single-stranded DNA in a sequence independent manner, protecting the DNA from cleavage by endonucleases. Binds the HipHop-HOAP complex to form terminin telomere-capping complex which binds to chromosome ends in a sequence-independent manner and prevents telomere fusion. (Adapted from ComplexPortal.)
    Protein Function (UniProtKB)
    Part of the HipHop-HOAP complex that recruits the MTV complex to form the terminin telomere-capping complex, which binds to chromosome ends in a sequence-independent manner and prevents telomere fusion (PubMed:11408576, PubMed:12510197, PubMed:12826664, PubMed:19181850, PubMed:19190990, PubMed:20057353). Telomere capping is independent of the origin recognition complex (ORC) (PubMed:12510197).
    (UniProt, Q95RV2)
    Summary (Interactive Fly)

    poorly conserved component of the telomere capping complex - prevents end fusion by maintaining a chromatin state that is independent of the underlying DNA sequence

    Gene Model and Products
    Number of Transcripts
    4
    Number of Unique Polypeptides
    3

    Please see the JBrowse view of Dmel\cav 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 Q95RV2)

    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 5.39

    Gene model reviewed during 5.46

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

    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    FBtr0084558
    1520
    345
    FBtr0084559
    1226
    337
    FBtr0084557
    1649
    337
    FBtr0084556
    1776
    343
    Additional Transcript Data and Comments
    Reported size (kB)
    Comments
    External Data
    Crossreferences
    Polypeptide Data
    Annotated Polypeptides
    Name
    FlyBase ID
    Predicted MW (kDa)
    Length (aa)
    Theoretical pI
    UniProt
    RefSeq ID
    GenBank
    FBpp0083943
    39.4
    345
    4.64
    FBpp0083944
    38.5
    337
    4.64
    FBpp0083942
    38.5
    337
    4.64
    FBpp0083941
    39.2
    343
    4.65
    Polypeptides with Identical Sequences

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

    337 aa isoforms: cav-PB, cav-PD
    Additional Polypeptide Data and Comments
    Reported size (kDa)
    Comments
    External Data
    Subunit Structure (UniProtKB)

    Component of the HipHop-HOAP telomere capping complex, composed of at least HipHop and cav/HOAP, and may include Su(var)205/HP1; HipHop and cav/HOAP, but not Su(var)205, are interdependent for their protein stability (PubMed:20057353). Interacts with HipHop (via N-terminus) (PubMed:20057353). Interacts (via C-terminus) with Su(var)205/HP1 dimer (via hinge and chromoshadow domain) and Orc1; possibly interacts with other components of the origin recognition complex (ORC) (PubMed:11408576, PubMed:12826664, PubMed:19181850). Each molecule of cav/HOAP interacts with 2 molecules of Su(var)205/HP1 (PubMed:12826664). The HipHop-HOAP complex recruits the MTV complex, consisting of moi/modigliani, tea and ver/verrocchio, to telomeres, forming the terminin telomere-capping complex (PubMed:19181850, PubMed:20679394). Interacts with moi/modigliani; the interaction is direct (PubMed:19181850). Interacts with ver/verrochio; the interaction is direct (PubMed:20679394). Interacts with HP6, which is also part of the terminin complex (PubMed:19190990). Interacts (via N-terminus) with peo/pendolino (via N-terminus); the interaction is direct (PubMed:26110638).

    (UniProt, Q95RV2)
    Crossreferences
    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\cav 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).

    0.59

    Transcript Expression
    Polypeptide Expression
    distribution deduced from reporter or direct label
    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\cav 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) 9-10
    • Stages(s) 11-12
    Alleles, Insertions, Transgenic Constructs, and Aberrations
    Classical and Insertion Alleles ( 13 )
    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 cav
    Transgenic constructs containing regulatory region of cav
    Aberrations (Deficiencies and Duplications) ( 2 )
    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
    metaphase & condensed nuclear chromosome
    Orthologs
    Human Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    Homo sapiens (Human) (0)
    Model Organism Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    Rattus norvegicus (Norway rat) (0)
    Mus musculus (laboratory mouse) (0)
    Xenopus tropicalis (Western clawed frog) (0)
    Danio rerio (Zebrafish) (0)
    Caenorhabditis elegans (Nematode, roundworm) (0)
    Anopheles gambiae (African malaria mosquito) (0)
    Arabidopsis thaliana (thale-cress) (0)
    Saccharomyces cerevisiae (Brewer's yeast) (0)
    Schizosaccharomyces pombe (Fission yeast) (0)
    Escherichia coli (enterobacterium) (0)
    Other Organism Orthologs (via OrthoDB)
    Data provided directly from OrthoDB:cav. Refer to their site for version information.
    Paralogs
    Paralogs (via DIOPT v9.1)
    Human Disease Associations
    FlyBase Human Disease Model Reports
      Disease Ontology (DO) Annotations
      Models Based on Experimental Evidence ( 0 )
      Allele
      Disease
      Evidence
      References
      Potential Models Based on Orthology ( 0 )
      Human Ortholog
      Disease
      Evidence
      References
      Modifiers Based on Experimental Evidence ( 0 )
      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.
      Homo sapiens (Human)
      Gene name
      Score
      OMIM
      OMIM Phenotype
      DO term
      Complementation?
      Transgene?
      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
      Interaction Browsers

      Please see the Physical Interaction reports below for full details
      protein-protein
      Physical Interaction
      Assay
      References
      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
      suppressible
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      External Data
      Subunit Structure (UniProtKB)
      Component of the HipHop-HOAP telomere capping complex, composed of at least HipHop and cav/HOAP, and may include Su(var)205/HP1; HipHop and cav/HOAP, but not Su(var)205, are interdependent for their protein stability (PubMed:20057353). Interacts with HipHop (via N-terminus) (PubMed:20057353). Interacts (via C-terminus) with Su(var)205/HP1 dimer (via hinge and chromoshadow domain) and Orc1; possibly interacts with other components of the origin recognition complex (ORC) (PubMed:11408576, PubMed:12826664, PubMed:19181850). Each molecule of cav/HOAP interacts with 2 molecules of Su(var)205/HP1 (PubMed:12826664). The HipHop-HOAP complex recruits the MTV complex, consisting of moi/modigliani, tea and ver/verrocchio, to telomeres, forming the terminin telomere-capping complex (PubMed:19181850, PubMed:20679394). Interacts with moi/modigliani; the interaction is direct (PubMed:19181850). Interacts with ver/verrochio; the interaction is direct (PubMed:20679394). Interacts with HP6, which is also part of the terminin complex (PubMed:19190990). Interacts (via N-terminus) with peo/pendolino (via N-terminus); the interaction is direct (PubMed:26110638).
      (UniProt, Q95RV2 )
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      DroID - A comprehensive database of gene and protein interactions.
      MIST (protein-protein) - An integrated Molecular Interaction Database
      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
      3-84
      Cytogenetic map
      Sequence location
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      95F2-95F2
      Limits computationally determined from genome sequence between P{PZ}Syx1A06737&P{PZ}Syx1A10660 and P{PZ}crb07207&P{PZ}BRWD305842
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      95D-95E
      (determined by in situ hybridisation)
      Experimentally Determined Recombination Data
      Location
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (10)
      Genomic Clones (14)
       

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

      cDNA Clones (85)
       

      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)
      BDGP DGC clones
      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

        cav protein binds specific heterochromatin satellite DNA sequences in gel mobility shift assays.

        Mutant larvae have increased frequencies of telomeric fusions in brain cells.

        Quantifying rates of protein sequence divergence within and between species reveals that the Drosophila genome harbors a substantial proportion of genes with a very high divergence rate.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: cav Hoap

        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: anon- EST:fe1G5 CG6219

        Source for identity of: Hoap CG6219

        Nomenclature comments
        Etymology

        The gene is named "caravaggio", because the linear multicentric chromosomes that characterize the mutant phenotype resemble little trains; there is an Italian train named "Caravaggio" in memory of the famous painter.

        Synonyms and Secondary IDs (23)
        Reported As
        Symbol Synonym
        Caravaggio
        anon-EST:fe1G5
        Secondary FlyBase IDs
        • FBgn0020360
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 77 )
        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 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
        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
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (160)