FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\twe
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General Information
Symbol
Dmel\twe
Species
D. melanogaster
Name
twine
Annotation Symbol
CG4965
Feature Type
FlyBase ID
FBgn0002673
Gene Model Status
Stock Availability
Enzyme Name (EC)
protein-tyrosine-phosphatase (3.1.3.48)
Gene Summary
twine (twe) encodes a putative tyrosine phosphatase. It regulates the meiotic cell cycle and contributes to spermatogenesis. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

Cdc25, twn, BG:DS02740.1 , l(2)35Fh, mat(2)synHB5

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-52
RefSeq locus
NT_033779 REGION:16259414..16261877
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (14 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 sequence or structural similarity
inferred from biological aspect of ancestor with PANTHER:PTN000850864
inferred from electronic annotation with InterPro:IPR000751
Biological Process (10 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
inferred from high throughput mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in spermatogenesis
inferred from mutant phenotype
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
located_in spindle
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN000850864
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000850864
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the MPI phosphatase family. (Q03019)
Catalytic Activity (EC/Rhea)
protein tyrosine phosphatase activity
O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate (3.1.3.48)
RHEA 10684:
Summaries
Gene Snapshot
twine (twe) encodes a putative tyrosine phosphatase. It regulates the meiotic cell cycle and contributes to spermatogenesis. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
CDC25 FAMILY PROTEIN PHOSPHATASES -
CDC25 phosphatases are tyrosine phosphatases that play critical roles in mitosis. The family is represented in Drosophila by the paralogous genes stg and twe. stg plays a role at the start of mitotic events during embryogenesis, oogenesis and regulates proliferation and stem cell maintenance, whereas twe plays a role in meiosis and oogenesis. (Adapted from FBrf0227974.)
Protein Function (UniProtKB)
Required during meiosis. Regulates the transition from the extended G2 phase to the onset of the first meiotic division.
(UniProt, Q03019)
Summary (Interactive Fly)

cdc25 phosphatase - dephosphorylation of cdc2 by Twine controls entry into meiosis in both males and females

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

Please see the JBrowse view of Dmel\twe 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 Q03019)

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

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

Gene model reviewed during 5.44

Gene model reviewed during 5.55

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0080855
2379
426
FBtr0331444
1803
426
FBtr0331445
2240
426
FBtr0331446
1916
426
Additional Transcript Data and Comments
Reported size (kB)

2 (northern blot)

1.738 (longest cDNA)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0080412
48.3
426
5.16
FBpp0303861
48.3
426
5.16
FBpp0303862
48.3
426
5.16
FBpp0303863
48.3
426
5.16
Polypeptides with Identical Sequences

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

426 aa isoforms: twe-PA, twe-PB, twe-PC, twe-PD
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\twe 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).

1.73

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

twe transcripts are expressed maximally in nurse cells in the later stages of oogenesis and the transcripts are transferred to the oocyte from stage 11 onward. The maternally derived transcripts are uniformly distributed in the cytoplasm of the early embryo and are excluded from the forming cells during cellularization. twe transcripts are also expressed in the male testis where they accumulate in the growing phase of spermatocytes. No zygotic twe transcription was detected.

twe expression is restricted to male and female gonads. In the ovary, twe is first detected in stage S7 nurse cells and is detected at high levels by stage S10. At stage S11, transcripts are translocated from the nurse cells to the oocyte. twe signal remains strong in early embryos up through cellularization. In the testis, no expression is observed in the apical tip where the mitotic divisions occur but rather in the premeiotic cyst of 16 primary spermatocytes and also after meiosis I in the 32-cell cysts.

Marker for
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
located_in spindle
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{twe-lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\twe 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 ( 7 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 33 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of twe
Transgenic constructs containing regulatory region of twe
Aberrations (Deficiencies and Duplications) ( 16 )
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
spermatocyte & meiotic cell cycle
spindle & oocyte
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (4)
12 of 14
Yes
Yes
1  
11 of 14
No
Yes
11 of 14
No
Yes
1  
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) (3)
12 of 14
Yes
Yes
11 of 14
No
Yes
10 of 14
No
Yes
Mus musculus (laboratory mouse) (3)
12 of 14
Yes
Yes
10 of 14
No
Yes
10 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (6)
8 of 13
Yes
Yes
7 of 13
No
Yes
6 of 13
No
Yes
4 of 13
No
Yes
3 of 13
No
Yes
1 of 13
No
No
Danio rerio (Zebrafish) (3)
12 of 14
Yes
Yes
5 of 14
No
Yes
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (4)
11 of 14
Yes
Yes
9 of 14
No
Yes
8 of 14
No
Yes
5 of 14
No
No
Anopheles gambiae (African malaria mosquito) (1)
Arabidopsis thaliana (thale-cress) (2)
2 of 13
Yes
Yes
1 of 13
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (3)
9 of 13
Yes
Yes
2 of 13
No
Yes
2 of 13
No
Yes
Schizosaccharomyces pombe (Fission yeast) (2)
8 of 12
Yes
Yes
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:twe. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (1)
11 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.
    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

    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
    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
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map
    2-52
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    35F1-35F1
    Limits computationally determined from genome sequence between P{lacW}CycEk05007&P{lacW}CycEk02602 and P{EP}crpEP721&P{lacW}crpk07829
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    35F-35F
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (15)
    Genomic Clones (27)
    cDNA Clones (78)
     

    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

        dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

        twe is required for both oogenesis and male meiosis and genetic evidence suggests that twe is a vital gene.

        da and crp are two independent DNA-binding components of Sebp3, a protein complex that appears to be required for the transcriptional activation of Sgs4. Since Sebp3 binding activity is specific for the salivary gland, and da and crp are ubiquitously expressed, the Sebp3 complex probably contains additional protein(s). In direction of increasing cytology: twe- crp- pkaap+

        Meiotic entry failure in bol mutants can be overcome by heterologous expression of twe.

        aly is required for CycB and twe expression in primary spermatocytes. Accumulation of twe and CycB proteins occurs just before the first meiotic division in wild type.

        "l(2)35Fh" gene is inferred from the lethality of In(2L)el18/Df(2L)RN2 - they should only overlap for twe but twe is not a lethal.

        stg and twe complement each other in the germ-line and early embryo. Lowering the maternal dose of stg and twe can advance the maternal/zygotic transition (MZT). Increasing twe, but not stg, can postpone the MZT.

        Mutations in can and sa are epistatic to mutations in twe.

        twe is not required for all aspects of the entry into male meiosis and mutations in twe lead to a variety of abnormal meiotic spindles and unusual chromosome segregation in female meiosis.

        twe is essential during oogenesis and has a role in regulating mitosis in the syncytial embryo stage.

        twe gene identified on basis of sequence similarity to cdc25 phosphatases by PCR with degenerate primers.

        Mutant embryos show defects occurring during syncytial blastoderm formation: cytoplasmic clearing appears irregular.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: twe l(2)35Fh

        Additional comments

        Can rescue allele 22 of Spom\cdc25, but not a deletion of the gene.

        Nomenclature History
        Source for database identify of

        Source for identity of: twe CG4965

        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (14)
        Reported As
        Symbol Synonym
        mat(2)syn-A
        mat(2)synHB5
        Secondary FlyBase IDs
        • FBgn0016862
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 68 )
        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
        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)
        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
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (215)