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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 Snapshot
twine (twe) encodes a putative tyrosine phosphatase. It regulates the meiotic cell cycle and contributes to spermatogenesis. [Date last reviewed: 2019-03-14]
Also Known As

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

Key Links
Genomic Location
Cytogenetic map
Sequence location
2L:16,259,414..16,261,877 [-]
Recombination map

2-52

RefSeq locus
NT_033779 REGION:16259414..16261877
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 (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:PTN000850864
(assigned by GO_Central )
inferred from sequence or structural similarity
Biological Process (12 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000089071
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000089071
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000089071
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000089071
(assigned by GO_Central )
inferred from electronic annotation with InterPro:IPR000751
(assigned by InterPro )
inferred from sequence or structural similarity
traceable author statement
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000850864
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000850864
(assigned by GO_Central )
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the MPI phosphatase family. (Q03019)
Catalytic Activity (EC)
Experimental Evidence
-
Predictions / Assertions
Protein tyrosine phosphate + H(2)O = protein tyrosine + phosphate (3.1.3.48)
Summaries
Gene Group (FlyBase)
DUAL SPECIFICITY PHOSPHATASES -
Dual Specificity Phosphatases (DSP) can dephosphorylate both tyrosine and serine/threonine residues. Additionally, DSPs may target a much larger and diverse set of substrates, including phosphoinositide lipids, RNA 5'-triphosphate and carbohydrates. (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
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

Sequence Ontology: Class of Gene
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
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
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
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
inferred from direct assay
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

GBrowse - Visual display of RNA-Seq signals

View Dmel\twe 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
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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 7 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 23 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of twe
Transgenic constructs containing regulatory region of twe
Deletions and Duplications ( 16 )
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 v8.0)
Homo sapiens (Human) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
11 of 15
Yes
No
10 of 15
No
Yes
10 of 15
No
Yes
1  
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
10 of 15
Yes
No
9 of 15
No
Yes
8 of 15
No
No
Rattus norvegicus (Norway rat) (3)
8 of 13
Yes
No
8 of 13
Yes
Yes
5 of 13
No
No
Xenopus tropicalis (Western clawed frog) (5)
8 of 12
Yes
No
8 of 12
Yes
No
6 of 12
No
Yes
2 of 12
No
Yes
2 of 12
No
Yes
Danio rerio (Zebrafish) (2)
10 of 15
Yes
No
7 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (4)
12 of 15
Yes
Yes
10 of 15
No
Yes
8 of 15
No
Yes
6 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (3)
10 of 15
Yes
Yes
2 of 15
No
Yes
1 of 15
No
Yes
Schizosaccharomyces pombe (Fission yeast) (2)
7 of 12
Yes
Yes
2 of 12
No
Yes
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG091908BI )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( None identified )
No non-Drosophilid orthologies identified
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W08P2 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0DFE )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0LFI )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (1)
8 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 ( 1 )
    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
    Interactions Browser
    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
    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 (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map

    2-52

    Cytogenetic map
    Sequence location
    2L:16,259,414..16,261,877 [-]
    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 (12)
    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 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: twe CG4965

    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.

    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.

    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 66 )
    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/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
    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
    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.
    modMine - A data warehouse for the modENCODE project
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DPiM - Drosophila Protein interaction map
    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
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    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
    References (189)