FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\ari-1
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General Information
Symbol
Dmel\ari-1
Species
D. melanogaster
Name
ariadne 1
Annotation Symbol
CG5659
Feature Type
FlyBase ID
FBgn0017418
Gene Model Status
Stock Availability
Enzyme Name (EC)
Gene Summary
ariadne 1 (ari-1) encodes an E3 ubiquitin ligase that interacts with the E2 ubiquitin conjugating enzyme encoded by Ubc10. The product of ari-1 and Ubc10 jointly ubiquitylate isoform A of the ecdysone receptor encoded by EcR. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
Also Known As

ari, ariadne, HL-V

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-59
RefSeq locus
NC_004354 REGION:18130392..18137784
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (16 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR018957
inferred from biological aspect of ancestor with PANTHER:PTN000188395
inferred from sequence model
inferred from electronic annotation with InterPro:IPR002867
Biological Process (7 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from mutant phenotype
involved_in oogenesis
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from direct assay
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR031127
Cellular Component (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
located_in nucleus
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN000188395
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000188512
inferred from biological aspect of ancestor with PANTHER:PTN000188395
Protein Family (UniProt)
Belongs to the RBR family. Ariadne subfamily. (Q94981)
Catalytic Activity (EC/Rhea)
ubiquitin protein ligase activity
Summaries
Gene Snapshot
ariadne 1 (ari-1) encodes an E3 ubiquitin ligase that interacts with the E2 ubiquitin conjugating enzyme encoded by Ubc10. The product of ari-1 and Ubc10 jointly ubiquitylate isoform A of the ecdysone receptor encoded by EcR. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
RING-BETWEEN-RING UBIQUITIN LIGASES -
E3 (ubiquitin) ligases interact with ubiquitin-E2 (ubiquitin-conjugating) enzymes and the target protein, transferring the ubiquitin from the E2 to the substrate. RING-between-RING (RBR) E3 ligases are characterised by three closely spaced domains: RING1 (a classical RING finger domain), a In-Betweeen-RING (IBR) domain and a RING2 domain. The mechanism of action is described as RING/HECT-like, with the RING1 domain facilitating E2-transfer (E3 RING-like) to form a thioester intermediate with a cysteine in RING2 which then modifies the substrate protein (HECT-like). (Adapted from PMID:24469331).
Protein Function (UniProtKB)
Atypical E3 ubiquitin-protein ligase, which catalyzes ubiquitination of target proteins together with ubiquitin-conjugating enzyme E2 Ubc10 (PubMed:10880484, PubMed:21900267, PubMed:29689197). Controls the subcellular localization and morphology of muscle nuclei (myonuclei) by regulating the protein levels and distribution of the LINC (LInker of Nucleoskeleton and Cytoskeleton) complex (PubMed:29689197). Functions by mediating the monoubiquitination of the LINC complex subunit koi leading to its subsequent proteasomal degradation (PubMed:29689197). Appears to function, at least partially redundantly, with the RBR E3 ligase family member park in nuclear localization and morphology (PubMed:29689197). Likely to function in metamorphosis by regulating the proteins levels of EcR isoform A (ECR-A) and its heterodimeric partner usp, via the ubiquitination and subsequent degradation of ECR-A (PubMed:21900267).
(UniProt, Q94981)
Summary (Interactive Fly)

E3 ubiquitin-ligase essential for neuronal development - mutants display a lower rate of spontaneous neurotransmitter release due to failures at the pre-synaptic side - evoked release in Ari-1 mutants was enhanced in a Ca(2+) dependent manner without modifications in the number of active zones, indicating that the probability of release per synapse is increased in these mutants - regulates myonuclear organization together with Parkin and is associated with aortic aneurysms

Gene Model and Products
Number of Transcripts
6
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\ari-1 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 Q94981)

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 5.53

Gene model reviewed during 5.46

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

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Gene model reviewed during 5.45

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Gene model reviewed during 5.56

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0089506
2656
503
FBtr0089507
2635
503
FBtr0089508
2811
503
FBtr0306057
4416
503
FBtr0332850
3234
503
FBtr0332851
3350
503
Additional Transcript Data and Comments
Reported size (kB)

4.3 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0088499
58.9
503
5.01
FBpp0088500
58.9
503
5.01
FBpp0089023
58.9
503
5.01
Polypeptides with Identical Sequences

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

Additional Polypeptide Data and Comments
Reported size (kDa)

503 (aa); 59 (kD observed); 59 (kD predicted)

Comments
External Data
Subunit Structure (UniProtKB)

Can form homodimers (PubMed:10880484, PubMed:29689197). Interacts (via RING-type 1 zinc finger) with Ubc10 (PubMed:10880484, PubMed:29689197). Interacts with the LINC complex member koi (PubMed:29689197). Interacts with park (PubMed:29689197). Interacts with ari-2 (PubMed:29689197). Specifically interacts with isoform ECR-A of EcR (PubMed:21900267).

(UniProt, Q94981)
Post Translational Modification

Autophosphorylated.

(UniProt, Q94981)
Domain

Members of the RBR family are atypical E3 ligases. They interact with E2 conjugating enzymes such as Ubc10 and function like HECT-type E3 enzymes: they bind E2s via the first RING-type zinc finger, but require an obligate trans-thiolation step during the ubiquitin transfer, requiring a conserved active site Cys residue in the second RING-type zinc finger. The active site probably forms a thioester intermediate with ubiquitin taken from the active-site cysteine of the E2 before ultimately transferring it to a Lys residue on the substrate.

(UniProt, Q94981)
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\ari-1 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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

ari-1 transcripts are detected at all developmental stages, with levels increasing from embryogenesis onwards. The peak is reached at metamorphosis and maintained in adults. ari-1 is detected in all tissues, with high levels in the adult CNS and female gonads.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Although ari-1 protein is detected mainly in the cytoplasm in Western blots of nuclear vs. cytoplasm enriched fractions, it does contain a putative nuclear localization signal.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
located_in nucleus
inferred from direct assay
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ari-1 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
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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 7 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ari-1
Transgenic constructs containing regulatory region of ari-1
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
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
photoreceptor cell R1 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R2 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R3 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R4 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R5 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R6 & rough endoplasmic reticulum | somatic clone
photoreceptor cell R7 & rough endoplasmic reticulum | somatic clone
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (16)
14 of 14
Yes
Yes
 
4  
5 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
12  
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (14)
14 of 14
Yes
Yes
5 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
Mus musculus (laboratory mouse) (15)
14 of 14
Yes
Yes
5 of 14
No
No
3 of 14
No
Yes
2 of 14
No
No
2 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (13)
11 of 13
Yes
Yes
3 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 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
Danio rerio (Zebrafish) (25)
14 of 14
Yes
Yes
14 of 14
Yes
Yes
5 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
2 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (14)
13 of 14
Yes
Yes
13 of 14
Yes
Yes
12 of 14
No
Yes
11 of 14
No
Yes
5 of 14
No
No
5 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (7)
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (45)
12 of 13
Yes
Yes
12 of 13
Yes
Yes
11 of 13
No
Yes
11 of 13
No
Yes
9 of 13
No
No
8 of 13
No
No
8 of 13
No
No
8 of 13
No
No
8 of 13
No
No
7 of 13
No
No
7 of 13
No
No
6 of 13
No
No
4 of 13
No
Yes
4 of 13
No
Yes
4 of 13
No
Yes
4 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
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
Yes
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (2)
12 of 13
Yes
Yes
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (2)
12 of 12
Yes
Yes
1 of 12
No
Yes
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:ari-1. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (6)
10 of 13
9 of 13
4 of 13
3 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 ( 0 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 1 )
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.
Dmel gene
Ortholog showing functional complementation
Supporting References
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)
Can form homodimers (PubMed:10880484, PubMed:29689197). Interacts (via RING-type 1 zinc finger) with Ubc10 (PubMed:10880484, PubMed:29689197). Interacts with the LINC complex member koi (PubMed:29689197). Interacts with park (PubMed:29689197). Interacts with ari-2 (PubMed:29689197). Specifically interacts with isoform ECR-A of EcR (PubMed:21900267).
(UniProt, Q94981 )
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
FlyBase
External Links
External Data
Linkouts
KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
X
Recombination map
1-59
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
16F7-16F7
Limits computationally determined from genome sequence between P{EP}EP970 and P{EP}ari-1EP317
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Experimentally Determined Recombination Data
Location

1-57.6--58.7

1-57.6-58.7

Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (35)
Genomic Clones (13)
 

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

cDNA Clones (124)
 

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

        ari-1 specifically binds the A isoform of EcR and ubiquitylates it.

        ari-1 is required for the correct differentiation of most cell types in the adult organism.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: ari l(1)16Fj

        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: ari CG5659

        Nomenclature comments
        Etymology

        The name "ariadne" refers to the loss of clues to find the correct path of axonal projection, and it evokes the Greek myth of the thread that Ariadne gave to Theseus to find his way in the Minotaur's labyrinth.

        Synonyms and Secondary IDs (13)
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 67 )
        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
        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
        FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        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
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        References (72)