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General Information
Symbol
Dmel\toc
Species
D. melanogaster
Name
toucan
Annotation Symbol
CG9660
Feature Type
FlyBase ID
FBgn0015600
Gene Model Status
Stock Availability
Gene Snapshot
toucan (toc) encodes a protein that associates with spindle microtubules being involved in mitotic spindle organization, mitotic sister chromatid segregation and ovarian follicle cell formation. [Date last reviewed: 2019-03-14]
Key Links
Genomic Location
Cytogenetic map
Sequence location
2L:3,068,345..3,144,786 [-]
Recombination map

2-8

RefSeq locus
NT_033779 REGION:3068345..3144786
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Molecular Function (GO)
    [Detailed GO annotations]
    Experimental Evidence
    -
    Predictions / Assertions
    Summaries
    Gene Model and Products
    Number of Transcripts
    10
    Number of Unique Polypeptides
    10

    Please see the GBrowse view of Dmel\toc or the JBrowse view of Dmel\toc 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_with_stop_codon_read_through ; SO:0000697

    Stop-codon suppression (UGA) postulated; FBrf0216884.

    Gene model reviewed during 5.44

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

    Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

    Gene model reviewed during 5.47

    Sequence Ontology: Class of Gene
    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    FBtr0077662
    8297
    2162
    FBtr0077665
    4854
    993
    FBtr0077664
    3121
    888
    FBtr0077663
    3410
    984
    FBtr0110903
    4787
    945
    FBtr0112883
    2525
    682
    FBtr0329906
    4265
    840
    FBtr0329907
    4787
    993
    FBtr0335388
    3266
    936
    FBtr0335389
    4628
    892
    Additional Transcript Data and Comments
    Reported size (kB)

    8, 7 (northern blot)

    Comments
    External Data
    Crossreferences
    Polypeptide Data
    Annotated Polypeptides
    Name
    FlyBase ID
    Predicted MW (kDa)
    Length (aa)
    Theoretical pI
    RefSeq ID
    GenBank
    FBpp0077346
    234.1
    2162
    9.06
    FBpp0077349
    109.9
    993
    7.13
    FBpp0077348
    98.5
    888
    6.90
    FBpp0077347
    108.2
    984
    7.96
    FBpp0110203
    104.6
    945
    7.89
    FBpp0111796
    76.6
    682
    5.65
    FBpp0302940
    93.2
    840
    5.16
    FBpp0302941
    110.0
    993
    6.05
    FBpp0307372
    103.1
    936
    7.07
    FBpp0307373
    99.3
    892
    7.05
    Polypeptides with Identical Sequences

    None of the polypeptides share 100% sequence identity.

    Additional Polypeptide Data and Comments
    Reported size (kDa)

    2176 (aa); 235 (kD predicted)

    Comments
    External Data
    Crossreferences
    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\toc using the Feature Mapper tool.

    External Data
    Crossreferences
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Linkouts
    Gene Ontology (11 terms)
    Molecular Function (1 term)
    Terms Based on Experimental Evidence (0 terms)
    Terms Based on Predictions or Assertions (1 term)
    CV Term
    Evidence
    References
    inferred from biological aspect of ancestor with PANTHER:PTN001271408
    (assigned by GO_Central )
    Biological Process (2 terms)
    Terms Based on Experimental Evidence (1 term)
    CV Term
    Evidence
    References
    Terms Based on Predictions or Assertions (1 term)
    CV Term
    Evidence
    References
    Cellular Component (8 terms)
    Terms Based on Experimental Evidence (7 terms)
    CV Term
    Evidence
    References
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    Terms Based on Predictions or Assertions (2 terms)
    CV Term
    Evidence
    References
    colocalizes_with cytoplasm
    inferred from biological aspect of ancestor with PANTHER:PTN001271408
    (assigned by GO_Central )
    inferred from biological aspect of ancestor with PANTHER:PTN001271408
    (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

    Comment: maternally deposited

    antennal anlage

    Comment: reported as procephalic ectoderm anlage

    central brain anlage

    Comment: reported as procephalic ectoderm anlage

    dorsal head epidermis anlage

    Comment: reported as procephalic ectoderm anlage

    visual anlage

    Comment: reported as procephalic ectoderm anlage

    ventral nerve cord primordium

    Comment: reported as ventral nerve cord anlage

    antennal primordium

    Comment: reported as procephalic ectoderm primordium

    central brain primordium

    Comment: reported as procephalic ectoderm primordium

    visual primordium

    Comment: reported as procephalic ectoderm primordium

    dorsal head epidermis primordium

    Comment: reported as procephalic ectoderm primordium

    lateral head epidermis primordium

    Comment: reported as procephalic ectoderm primordium

    ventral head epidermis primordium

    Comment: reported as procephalic ectoderm primordium

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

    Seven and 8 kb transcripts were detected in Northern blots of adults.

    toc transcript is detected in the germarium as soon as the 16-cell cysts are formed, and through stages S1-S2 of oogenesis. In stages S10A-S10B expression is detected in the nurse cell and in the posterior of the oocyte. Expression is uniform in the late stage oocyte and the early embryo. The expression pattern detected in a Ecol\lacZ enhancer trap (Ecol\lacZtoc-P) is consistent with toc transcript expression.

    Marker for
     
    Subcellular Localization
    CV Term
    Polypeptide Expression
    immunolocalization
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    organism
    nucleus

    Comment: absent by cycle 14

    organism

    Comment: telophase

    organism
    spindle

    Comment: prophase and metaphase

    organism
    midbody

    Comment: telophase

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

    toc protein is visible during all syncytial divisions in the embryo. Protein levels decrease during the 13th division,and are undetectable by embryonic cycle 14. Localization of toc protein is dynamic during the mitotic cycle. During interphase, toc localizes to the centrosomes and nuclear envelope. During prophase, toc protein is associated with the two half-spindles. At metaphase, toc is associated with the entire spindle. During anaphase, toc protein accumulates at the spindle poles, under the centrosomes. During telophase, toc protein accumulates at the midbody and in the centrosome area.

    Marker for
     
    Subcellular Localization
    CV Term
    Evidence
    References
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    inferred from direct assay
    Expression Deduced from Reporters
    High-Throughput Expression Data
    Associated Tools

    GBrowse - Visual display of RNA-Seq signals

    View Dmel\toc 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
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    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, and Transgenic Constructs
    Classical and Insertion Alleles ( 39 )
    For All Classical and Insertion Alleles Show
     
    Other relevant insertions
    Transgenic Constructs ( 3 )
    For All Alleles Carried on Transgenic Constructs Show
    Transgenic constructs containing/affecting coding region of toc
    Transgenic constructs containing regulatory region of toc
    Deletions and Duplications ( 28 )
    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 v7.1)
    Homo sapiens (Human) (1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    1 of 15
    Yes
    Yes
    Model Organism Orthologs (via DIOPT v7.1)
    Mus musculus (laboratory mouse) (1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    2 of 15
    Yes
    Yes
    Rattus norvegicus (Norway rat) (0)
    No records found.
    Xenopus tropicalis (Western clawed frog) (0)
    No records found.
    Danio rerio (Zebrafish) (1)
    1 of 15
    Yes
    Yes
    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.
    Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091900EU )
    Organism
    Common Name
    Gene
    AAA Syntenic Ortholog
    Multiple Dmel Genes in this Orthologous Group
    Drosophila melanogaster
    fruit fly
    Drosophila suzukii
    Spotted wing Drosophila
    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 willistoni
    Drosophila virilis
    Drosophila virilis
    Drosophila mojavensis
    Drosophila grimshawi
    Drosophila grimshawi
    Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091501R6 )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Musca domestica
    House fly
    Glossina morsitans
    Tsetse fly
    Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
    No non-Dipteran orthologies identified
    Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
    No non-Insect Arthropod orthologies identified
    Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G02AW )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Ciona intestinalis
    Vase tunicate
    Paralogs
    Paralogs (via DIOPT v7.1)
    Drosophila melanogaster (Fruit fly) (0)
    No records found.
    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 v7.1 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
      esyN Network Diagram
      Interactions Browser
      Summary of Genetic Interactions
      esyN Network Diagram
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      External Data
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      DroID - A comprehensive database of gene and protein interactions.
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      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)
      2L
      Recombination map

      2-8

      Cytogenetic map
      Sequence location
      2L:3,068,345..3,144,786 [-]
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      23D1-23D2
      Limits computationally determined from genome sequence between P{PZ}lilli00632 and P{PZ}toc01361
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      23D1-23D2
      (determined by in situ hybridisation)
      23D-23D
      (determined by in situ hybridisation)
      Experimentally Determined Recombination Data
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (22)
      Genomic Clones (71)
      cDNA Clones (133)
       

      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
      DRSC - Results frm RNAi screens
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      Antibody Information
      Laboratory Generated Antibodies
       

      polyclonal

      Commercially Available Antibodies
       
      Other Information
      Relationship to Other Genes
      Source for database identify of
      Source for database merge of

      Source for merge of: toc BcDNA:LD27161

      Additional comments

      Source for merge of toc BcDNA:LD27161 was a shared cDNA ( date:020730 ).

      Other Comments

      Candidate stable intronic sequence RNA (sisRNA) identified within 5'UTR of this gene.

      toc is essential for the assembly and function of the syncytial mitotic spindles.

      Dominant interactions indicate that toc is acting in the same signalling pathways for the formation of the egg chamber as da, N and Dl.

      toc is expressed and required in germline cells to promote proper differentiation of the somatic follicle cells. Results propose toc participates in the production or localisation of a germline-specific signal(s) that is required for the patterning of the follicular epithelium.

      Mutant ovaries are defective in the enclosure of newly formed germline cysts by the follicle cells, the formation of interfollicular stalks, the migration of follicle cells over the oocyte and the formation of the eggshell. Overexpression of toc causes the production of longer interfollicular stalks.

      Identified via an enhancer-trap line showing maternal effect female sterility.

      Analysis of revertant alleles of tocP suggest toc is required for communciation between germline cells and somatic cells during oogenesis.

      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 92 )
      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/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
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      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
      KEGG Genes - Molecular building blocks of life in the genomic space.
      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
      FLIGHT - Cell culture data for RNAi and other high-throughput technologies
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      FlyMine - An integrated database for Drosophila genomics
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (10)
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (86)