FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\toc
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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 Summary
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] (FlyBase Gene Snapshot)
Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-8
RefSeq locus
NT_033779 REGION:3068345..3144786
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (10 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
Biological Process (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (8 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
located_in cell cortex
inferred from direct assay
located_in centrosome
inferred from direct assay
located_in midbody
inferred from direct assay
located_in nuclear membrane
inferred from direct assay
located_in nucleus
inferred from direct assay
located_in spindle pole
inferred from direct assay
located_in spindle
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
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN001271408
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
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]
Gene Model and Products
Number of Transcripts
10
Number of Unique Polypeptides
10

Please see 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
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VQM0)

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

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

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
UniProt
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
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
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).

4.15

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 EcollacZ enhancer trap (EcollacZtoc-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
located_in cell cortex
inferred from direct assay
located_in centrosome
inferred from direct assay
located_in midbody
inferred from direct assay
located_in nuclear membrane
inferred from direct assay
located_in nucleus
inferred from direct assay
located_in spindle pole
inferred from direct assay
located_in spindle
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\toc 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 ( 42 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 5 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of toc
Transgenic constructs containing regulatory region of toc
Aberrations (Deficiencies and Duplications) ( 29 )
Inferred from experimentation ( 29 )
Inferred from location ( 10 )
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) (6)
3 of 14
Yes
Yes
3 of 14
Yes
Yes
2 of 14
No
Yes
1 of 14
No
Yes
1 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) (5)
3 of 14
Yes
Yes
3 of 14
Yes
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (5)
5 of 14
Yes
Yes
3 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (5)
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
No
Danio rerio (Zebrafish) (11)
3 of 14
Yes
Yes
3 of 14
Yes
Yes
3 of 14
Yes
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
1 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (3)
3 of 12
Yes
Yes
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:toc. 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
    Summary of Genetic Interactions
    Interaction Browsers
    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.
    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)
    2L
    Recombination map
    2-8
    Cytogenetic map
    Sequence location
    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 (63)
    Genomic Clones (71)
    cDNA Clones (49)
     

    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
     

    polyclonal

    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
        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.

        Relationship to Other Genes
        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 ).

        Nomenclature History
        Source for database identify of
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (12)
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 97 )
        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/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)
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (91)