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General Information
Symbol
Dmel\exu
Species
D. melanogaster
Name
exuperantia
Annotation Symbol
CG8994
Feature Type
FlyBase ID
FBgn0000615
Gene Model Status
Stock Availability
Gene Summary
Ensures the proper localization of the mRNA of the bicoid gene to the anterior regions of the oocyte thus playing a fundamental role in the establishment of the polarity of the oocyte. May bind the bcd mRNA. (UniProt, P28750)
Contribute a Gene Snapshot for this gene.
Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-91
RefSeq locus
NT_033778 REGION:20667427..20671083
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (16 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from physical interaction with UniProtKB:P23128
(assigned by UniProt )
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from key residues
Biological Process (8 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from expression pattern
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from high throughput mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
Cellular Component (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
located_in cytoplasm
inferred from direct assay
(assigned by UniProt )
located_in P-body
inferred from direct assay
inferred from high throughput direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
located_in cytoplasm
traceable author statement
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
Protein Function (UniProtKB)
Ensures the proper localization of the mRNA of the bicoid gene to the anterior regions of the oocyte thus playing a fundamental role in the establishment of the polarity of the oocyte. May bind the bcd mRNA.
(UniProt, P28750)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
exu: exuperantia (T. Schupbach)
Maternal-effect lethal mutant; embryos from homozygous mothers lack anterior-most head structures. Instead they form an inverted posterior midgut and proctodeal region at their anterior end. Cephalic furrow at gastrulation is shifted towards anterior. Analysis of germline clones indicates that the mutation is germline autonomous (Schupbach and Wieschaus, l986, Dev. Biol. 113: 443-48). Causes shift in blastoderm fate map as indicated by ftz expression; thoracic stripes broadened and shifted anteriorly; abdominal stripes narrowed and compressed posteriorly (Mlodzik et al.). Eggs produced by exu mothers appear to have a more uniform distribution of bcd+ product, i.e., less concentration anteriorly (Fronhofer and Nusslein-Volhard).
Summary (Interactive Fly)

a noncanonical RNA-binding protein with EXO-SAM-like domain architecture that interacts with its target RNA as a homodimer a core component of a large protein complex involved in localizing mRNAs both within nurse cells and the developing oocyte

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\exu 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

Gene model reviewed during 5.51

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0086242
2320
532
FBtr0086243
1988
532
FBtr0086244
2621
532
FBtr0302285
2631
532
Additional Transcript Data and Comments
Reported size (kB)

3.0, 2.3 (northern blot)

2.9, 2.5, 2.1 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0085555
58.0
532
10.11
FBpp0085556
58.0
532
10.11
FBpp0085557
58.0
532
10.11
FBpp0291491
58.0
532
10.11
Polypeptides with Identical Sequences

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

532 aa isoforms: exu-PA, exu-PB, exu-PC, exu-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

532 (aa); 58 (kD)

Comments
External Data
Subunit Structure (UniProtKB)

Component of the osk RNP complex, which is composed of at least exuperantia (exu), ypsilon schachtel (yps), aret (bruno), cup, and the mRNA of osk (PubMed:10662770). In the sponge body, forms a ribonucleoprotein complex (RNP) containing at least me31B, exu, yps and the mRNA of osk; interactions with exu and yps are RNA dependent (PubMed:11546740).

(UniProt, P28750)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\exu 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
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
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
pole plasm

Comment: maternally deposited

organism

Comment: maternally deposited

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

The male exu transcript is present in late larval, pupal, and adult stages.

The exu 2.9kb transcript is specific to males and is limited to the germline, since flies without germ cells, produced from tud mothers, lack sex-specific transcripts.

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
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
located_in cytoplasm
inferred from direct assay
(assigned by UniProt )
located_in P-body
inferred from direct assay
inferred from high throughput direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\exu 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
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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 31 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of exu
Transgenic constructs containing regulatory region of exu
Aberrations (Deficiencies and Duplications) ( 9 )
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 v8.0)
Homo sapiens (Human) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1  
1 of 15
Yes
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
No
Rattus norvegicus (Norway rat) (2)
1 of 13
Yes
No
1 of 13
Yes
No
Xenopus tropicalis (Western clawed frog) (1)
1 of 12
Yes
Yes
Danio rerio (Zebrafish) (2)
Caenorhabditis elegans (Nematode, roundworm) (0)
No records found.
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG0919078J )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091509AZ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0AB0 )
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
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0A7B )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (1)
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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 v8.0 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
    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
    RNA-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)
    Component of the osk RNP complex, which is composed of at least exuperantia (exu), ypsilon schachtel (yps), aret (bruno), cup, and the mRNA of osk (PubMed:10662770). In the sponge body, forms a ribonucleoprotein complex (RNP) containing at least me31B, exu, yps and the mRNA of osk; interactions with exu and yps are RNA dependent (PubMed:11546740).
    (UniProt, P28750 )
    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
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-91
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    57A9-57A10
    Limits computationally determined from genome sequence between P{lacW}l(2)k02206k02206 and P{lacW}l(2)k06409k06409&P{PZ}insc05475
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    57A4-57B1
    (determined by in situ hybridisation)
    Tentative cytological location determined by segmental aneuploidy.
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (22)
    Genomic Clones (26)
    cDNA Clones (154)
     

    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 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
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of

    Source for identity of: exu CG8994

    Source for database merge of
    Additional comments
    Other Comments

    exu protein is highly enriched in the sponge bodies, subcellular structures in nurse cells. Neither the accumulation of, or level of exu protein is dependent on the amount of bcd mRNA present in the ovaries. Results propose that sponge bodies are structures that, by assembly and transport of included molecules or associated structures, are involved in localisation of mRNAs in oocytes.

    Deletions within the male-specific 3'-UTR lead to male sterility associated with reduced steady-state levels of the mutant mRNA in vivo.

    An ovarian protein, exl binds to BLE1, an RNA localisation element from bcd mRNA. exl localisation is disrupted in exu mutants suggesting exl and exu proteins interact.

    tra2 is required in male germ cells for efficient male-specific processing of exu RNA. In the absence of tra2 males produce a new exu mRNA which is processed at its 3' end so that it contains sequences normally specific to the female 3' untranslated region. The male and female mRNAs differ in the untranslated regions so the predicted polypeptide is the same.

    Drug studies indicate that the Avic\GFP exu fusion protein is co-localised with microtubules during oogenesis. These particles are RNP complexes which transport and target bcd mRNA to the oocyte anterior cortex.

    In vitro binding and precipitation assays show that bacterially expressed exu protein binds to tubulin. Particles seen with the exu Avic\GFP fusion protein are RNP complexes that bind bcd mRNA, the particles associate with microtubules via exu in nurse cells and is transported along the microtubules and targetted to the anterior cortex of the oocyte.

    Comparison of bcd and hts transcript distribution in swa, exu and stau mutant embryos indicates different genetic requirements for proper localization of bcd and hts RNAs.

    exu is required for the proper localization of bcd mRNA in the oocyte.

    bcd mRNA localization requires the function of at least three genes, the exu gene acts earliest in the pathway. exu acts in initiating bcd mRNA localization but does not play a persistant role in that process.

    exu is not required for the maintenance of bcd mRNA localization during late stages of oogenesis or early embryogenesis. It is required for the establishment of bcd mRNA localization in the developing oocyte.

    Mutations in maternal anterior class gene exu interact with RpII140wimp.

    The effects of taxol treatment on exu mutants suggest general roles for the exu gene product in the multi-step bcd mRNA localization process.

    In embryos from exu mutant females bcd transcripts are delocalized and exposed to nos activity in the posterior.

    exu mutants exhibit weak anterior deletions, and weak segmentation defects in the posterior abdomen.

    Mature follicles are immunologically stained for asymmetric distribution of ecdysteroid-related antigen. During late oogenesis localisation of the antigen changes dramatically suggesting the antigen plays a role in early embryogenesis and, perhaps, in pattern formation.

    Mutations in exu result in a maternal effect phenotype with defects during the early stages of gastrulation and defects in the anteroposterior axis.

    exu mutations disrupt the second stage of bcd RNA localization during oogenesis, localization to the apical regions of the nurse cells.

    Mutations at exu cause anterior defects, rarely posterior defects in the embryo.

    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 73 )
    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/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
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    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
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    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
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    InterPro - A database of protein families, domains and functional sites
    KEGG Genes - Molecular building blocks of life in the genomic space.
    MARRVEL_MODEL - MARRVEL (model organism gene)
    modMine - A data warehouse for the modENCODE project
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
    FlyMine - An integrated database for Drosophila genomics
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (8)
    Reported As
    Secondary FlyBase IDs
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (250)