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General Information
Symbol
Dmel\tsr
Species
D. melanogaster
Name
twinstar
Annotation Symbol
CG4254
Feature Type
FlyBase ID
FBgn0011726
Gene Model Status
Stock Availability
Gene Summary
Exhibits F-actin depolymerizing activity and regulates actin cytoskeleton dynamics (PubMed:21205790). Required for cytokinesis in both mitotic and meiotic cells and for aster migration and separation (PubMed:8522587). Promotes cell motility during ovary development and oogenesis (PubMed:11175754). During larval development, required for the cell rearrangement needed for formation of terminal filaments which are stacks of somatic cells that are important for the initiation of ovarioles (PubMed:11175754). Also required for border cell migration during oogenesis (PubMed:11175754). During border cell migration, required for actin turnover and lamellipodial protrusion (PubMed:21205790). Required for the establishment of planar cell polarity (PCP) where cells adopt a uniform orientation within the plane of an epithelium (PubMed:16571634). During establishment of PCP, required for the redistribution of the PCP core proteins fz and stan/fmi to the proximodistal cell boundary (PubMed:16571634). During pupal development, required for elongation of the retinal cell body and for rhabdomere morphogenesis (PubMed:18423434). Required for mushroom body neuroblast proliferation and axon growth (PubMed:15572110). Plays a role in the positive regulation of protein secretion (PubMed:20026655). Plays a role in the regulation of nuclear localization of actin (PubMed:22323606). Required for the maintenance of epithelial integrity by controlling cell junctions and is also necessary for cell survival and tissue growth through regulation of JNK and yki signaling (PubMed:27041568). (UniProt, P45594)
Contribute a Gene Snapshot for this gene.
Also Known As

cofilin, ntf, l(2)k05633, D-cof, ADF

Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:24,044,453..24,046,805 [-]
Recombination map
2-106
RefSeq locus
NT_033778 REGION:24044453..24046805
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (32 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:1874
inferred from sequence or structural similarity with SGD:S000003973
inferred from biological aspect of ancestor with PANTHER:PTN000227258
(assigned by GO_Central )
Biological Process (26 terms)
Terms Based on Experimental Evidence (22 terms)
CV Term
Evidence
References
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
involved_in axonogenesis
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
involved_in cell morphogenesis
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from genetic interaction with FLYBASE:mbt; FB:FBgn0025743
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000227258
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000227261
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000227259
(assigned by GO_Central )
involved_in cell motility
inferred from biological aspect of ancestor with PANTHER:PTN000227259
(assigned by GO_Central )
Cellular Component (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
located_in nucleoplasm
inferred from high throughput direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in actin cytoskeleton
inferred from biological aspect of ancestor with PANTHER:PTN000227258
(assigned by GO_Central )
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN000227258
(assigned by GO_Central )
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the actin-binding proteins ADF family. (P45594)
Summaries
Protein Function (UniProtKB)
Exhibits F-actin depolymerizing activity and regulates actin cytoskeleton dynamics (PubMed:21205790). Required for cytokinesis in both mitotic and meiotic cells and for aster migration and separation (PubMed:8522587). Promotes cell motility during ovary development and oogenesis (PubMed:11175754). During larval development, required for the cell rearrangement needed for formation of terminal filaments which are stacks of somatic cells that are important for the initiation of ovarioles (PubMed:11175754). Also required for border cell migration during oogenesis (PubMed:11175754). During border cell migration, required for actin turnover and lamellipodial protrusion (PubMed:21205790). Required for the establishment of planar cell polarity (PCP) where cells adopt a uniform orientation within the plane of an epithelium (PubMed:16571634). During establishment of PCP, required for the redistribution of the PCP core proteins fz and stan/fmi to the proximodistal cell boundary (PubMed:16571634). During pupal development, required for elongation of the retinal cell body and for rhabdomere morphogenesis (PubMed:18423434). Required for mushroom body neuroblast proliferation and axon growth (PubMed:15572110). Plays a role in the positive regulation of protein secretion (PubMed:20026655). Plays a role in the regulation of nuclear localization of actin (PubMed:22323606). Required for the maintenance of epithelial integrity by controlling cell junctions and is also necessary for cell survival and tissue growth through regulation of JNK and yki signaling (PubMed:27041568).
(UniProt, P45594)
Summary (Interactive Fly)

Drosophila cofilin - actin-depolymerizing and actin-severing protein - essential for neuronal axon growth

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

Please see the JBrowse view of Dmel\tsr 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

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

Gene model reviewed during 5.52

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0072188
735
148
Additional Transcript Data and Comments
Reported size (kB)

0.8-0.9 (northern blot)

0.7 (longest cDNA)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0072097
17.2
148
7.24
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)

17 (aa); 148 (kD)

Comments
External Data
Post Translational Modification

Phosphorylated in vitro by protein kinase LIMK1 (PubMed:20026655). Phosphorylation is required for inactivation of tsr and for cell proliferation and axon growth (PubMed:15572110). Phosphorylation is negatively regulated by the panthothenate kinase fbl which catalyzes the first step in the conversion of panthothenic acid to coenzyme A (PubMed:22912811).

Dephosphorylated by protein phosphatase ssh which activates tsr.

(UniProt, P45594)
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\tsr 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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

tsr transcript is detected throughout development, with peaks of expression in late larval and pupal stages.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

In stage S9,10 egg chambers, total tsr staining is strong in nurse cells and follicle epithelium, while phosphorylated tsr is strong in nurse cell cytoplasm and ring canals but weak in the follicular epithelium. Both are present in border cells but in distinct patterns. Total tsr is present in an asymmetric pattern with more staining at the leading edge while phosphorylated tsr staining is uniform.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
located_in nucleoplasm
inferred from high throughput direct assay
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\tsr 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
EMBL-EBI Single Cell Expression Atlas
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 37 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 21 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of tsr
Transgenic constructs containing regulatory region of tsr
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
actin cytoskeleton & eye disc | somatic clone
adult mushroom body & axon | somatic clone
aster & primary spermatocyte cyst
aster & secondary spermatocyte
follicle cell & actin filament | ectopic | somatic clone
nurse cell & nucleus (with tsr1)
nurse cell & nucleus (with tsrntf)
oocyte & actin filament | ectopic | germ-line clone
wing & filamentous actin, with tsr139.cBa, tsrV27Q.cBa
wing & filamentous actin, with tsrΔ96, tsr139.cBa
wing & filamentous actin, with tsrΔ96, tsrV27Q.cBa
wing & filamentous actin (with tsrV27Q.cBa)
wing & filamentous actin (with tsrΔ96)
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
Yes
 
2  
10 of 15
No
Yes
2  
9 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?
11 of 15
Yes
Yes
11 of 15
Yes
Yes
9 of 15
No
Yes
Rattus norvegicus (Norway rat) (4)
9 of 13
Yes
Yes
8 of 13
No
Yes
7 of 13
No
Yes
2 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
9 of 12
Yes
Yes
9 of 12
Yes
Yes
8 of 12
No
Yes
Danio rerio (Zebrafish) (3)
11 of 15
Yes
Yes
11 of 15
Yes
Yes
9 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (1)
12 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (12)
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
8 of 9
Yes
Yes
7 of 9
No
Yes
7 of 9
No
Yes
7 of 9
No
Yes
6 of 9
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
12 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (1)
10 of 12
Yes
Yes
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG09190HMV )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150DL6 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0GKV )
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
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0JU5 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0TDH )
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)
9 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 ( 1 )
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.
Dmel gene
Ortholog showing functional complementation
Supporting References
Interactions
Summary of Physical Interactions
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
Signaling Pathways (FlyBase)
Metabolic Pathways
External Data
Linkouts
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2R
Recombination map
2-106
Cytogenetic map
Sequence location
2R:24,044,453..24,046,805 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
60B5-60B5
Limits computationally determined from genome sequence between P{lacW}Phmk07623&P{lacW}tsrk05633 and P{EP}EP503
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
60B4-60B5
(determined by in situ hybridisation)
60A10-60B1
(determined by in situ hybridisation)
Also placed in 60A8--60A16 by deficiency mapping.
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (29)
Genomic Clones (13)
 

Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

cDNA Clones (324)
 

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)
    Other clones
    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: tsr CG4254

    Source for database merge of

    Source for merge of: tsr ntf

    Source for merge of: tsr l(2)k05633

    Additional comments
    Other Comments

    dsRNA made from templates generated with primers directed against this gene was tested in an RNAi screen to identify genes that regulate Ca2+ release.

    dsRNA directed against this gene causes defects in cytokinesis when tested in an RNAi screen in S2 cells.

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

    RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a phenotype when assayed in Kc167 and S2R+ cells: binucleate cells with increased or polarized (uneven) accumulation of F-actin.

    dsRNA made from templates generated with primers directed against this gene is tested in an RNAi screen for effects on actin-based lamella formation.

    tsr regulates actin filament formation throughout the cell cortex in the follicular epithelium.

    tsr is required for the formation of terminal filaments during larval development and is also required for the migration of border cells during oogenesis.

    tsr mutants show disruption of terminal filament (TF) formation in the ovary. TF precursor cells can be seen scattered in a ring in the anterior region of late third larval instar tsr mutant ovaries. tsr is pupal lethal.

    tsr is required for ovarian morphogenesis.

    Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.

    tsr mutant spermatocytes exhibit abnormal positioning and delayed migration of asters to cell poles.

    tsr has been cloned and characterised.

    Induced expression of tsr proteins in yeast causes aberrant cell shapes reflecting defects in cytokinesis and/or cell shape maintenance.

    tsr mutant spermatocytes exhibit abnormal positioning and delayed migration of asters to cell poles. tsr mutants also display a defect in cytokinesis (in mitosis and meiosis).

    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 41 )
    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
    Other crossreferences
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    EMBL-EBI Single Cell Expression Atlas
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    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.
    MARRVEL_MODEL
    modMine - A data warehouse for the modENCODE project
    Linkouts
    ApoDroso - Functional genomic database for photoreceptor development, survival and function
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    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 (31)
    Reported As
    Symbol Synonym
    ADF/cofilin
    Cadf
    cofilin/ADF
    cofilin/actin depolymerizing factor homolog
    l(2)k03101
    l(2)k13409
    Name Synonyms
    Cofilin/actin depolymerizing factor-like
    actin-depolymerizing factor
    no terminal filament
    Secondary FlyBase IDs
    • FBgn0011567
    • FBgn0020533
    • FBgn0020539
    • FBgn0022163
    • FBgn0024209
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (249)