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General Information
Symbol
Dmel\Tlk
Species
D. melanogaster
Name
Tousled-like kinase
Annotation Symbol
CG34412
Feature Type
FlyBase ID
FBgn0283657
Gene Model Status
Stock Availability
Gene Summary
Tousled-like kinase (Tlk) encodes a serine/threonine kinase that acts in a cell cycle-dependent manner. It contributes to cell morphogenesis, cell migration and cell death through interactions with chromatin components. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

l(1)G0054, BEST:GH17330 , BcDNA:GH07910 , EP1413

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-4
RefSeq locus
NC_004354 REGION:3720007..3789770
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (17 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:asf1; FB:FBgn0029094
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables ATP binding
inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR002290, InterPro:IPR017441
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN000540737
(assigned by GO_Central )
inferred from sequence or structural similarity with HGNC:11841
inferred from sequence or structural similarity with HGNC:11842
Biological Process (10 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:asf1; FB:FBgn0029094
inferred from genetic interaction with FLYBASE:mars; FB:FBgn0033845
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from direct assay
inferred from mutant phenotype
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000540737
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000540738
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000540737
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000540738
(assigned by GO_Central )
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in heterochromatin
inferred from direct assay
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000540737
(assigned by GO_Central )
Protein Family (UniProt)
-
Summaries
Gene Snapshot
Tousled-like kinase (Tlk) encodes a serine/threonine kinase that acts in a cell cycle-dependent manner. It contributes to cell morphogenesis, cell migration and cell death through interactions with chromatin components. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
OTHER CONVENTIONAL PROTEIN KINASE DOMAIN KINASES -
The group of 'other conventional protein kinase domain kinases' consists of kinases with a conventional eukaryotic protein kinase domain that do not fit into any of the other major groups. This grouping is described in FBrf0201870.
Gene Model and Products
Number of Transcripts
13
Number of Unique Polypeptides
9

Please see the JBrowse view of Dmel\Tlk 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 model reviewed during 5.55

3' UTR contains an ORF that is conserved among close Drosophila species (FBrf0202906; potential peptide 129 aa in length).

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

Due to significant sequence change in this region between Releases 4 and 5, data that predate Release 5 are not shown for some categories of evidence.

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

Gene model reviewed during 5.46

Stop-codon suppression (UAG) postulated; FBrf0243886.

Gene model reviewed during 6.32

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0112680
4624
1463
FBtr0070596
5849
1254
FBtr0299579
5537
1255
FBtr0299580
6513
1266
FBtr0299581
7243
1489
FBtr0299582
5890
1266
FBtr0299583
4673
1242
FBtr0301659
4882
1266
FBtr0333751
7403
1265
FBtr0333752
4308
1011
FBtr0333753
7490
1242
FBtr0345319
4355
1242
FBtr0479803
5890
1276
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0111592
160.3
1463
9.10
FBpp0099673
135.5
1254
9.31
FBpp0288854
135.6
1255
9.36
FBpp0288855
136.7
1266
9.10
FBpp0288856
162.2
1489
9.62
FBpp0288857
136.7
1266
9.10
FBpp0288858
133.9
1242
9.02
FBpp0290873
136.7
1266
9.10
FBpp0305893
136.4
1265
9.25
FBpp0305894
110.0
1011
9.50
FBpp0305895
133.9
1242
9.02
FBpp0311479
133.9
1242
9.02
FBpp0428136
137.8
1276
9.24
Polypeptides with Identical Sequences

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

1266 aa isoforms: Tlk-PF, Tlk-PH, Tlk-PJ
1242 aa isoforms: Tlk-PI, Tlk-PM, Tlk-PN
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
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\Tlk 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
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
nurse cell & oocyte

Comment: maternal transcript transported from nurse cells to oocytes

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

tlk is ubiquituously distributed in all proliferative tissues.

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

tlk protein is localized to the nucleus in interphase cells, and diffusely distributed throughout the cell during other parts of the mitotic cell cycle. tlk protein does not associate directly with DNA.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in heterochromatin
inferred from direct assay
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Tlk 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
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
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 83 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 20 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Tlk
Transgenic constructs containing regulatory region of Tlk
Aberrations (Deficiencies and Duplications) ( 0 )
Inferred from experimentation ( 0 )
Inferred from location ( 0 )
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
larval salivary gland & nuclear chromosome, with Scer\GAL4hs.2sev
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
11 of 15
Yes
Yes
8 of 15
No
Yes
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
11 of 15
Yes
Yes
8 of 15
No
Yes
Rattus norvegicus (Norway rat) (3)
7 of 13
Yes
Yes
5 of 13
No
Yes
4 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
9 of 12
Yes
Yes
4 of 12
No
Yes
1 of 12
No
Yes
Danio rerio (Zebrafish) (4)
10 of 15
Yes
Yes
7 of 15
No
Yes
6 of 15
No
Yes
3 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (2)
13 of 15
Yes
Yes
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (1)
5 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (5)
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG0919031Y )
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) ( EOG091502G5 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse 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) ( EOG090W02AZ )
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
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X028H )
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
Stegodyphus mimosarum
African social velvet spider
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) ( EOG091G02XG )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (1)
2 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 ( 0 )
    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
    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
    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)
    X
    Recombination map
    1-4
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    3E6-3E7
    Left limit from in situ hybridisation (FBrf0128367) Right limit from in situ hybridisation (FBrf0128367)
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    3F4-3F5
    (determined by in situ hybridisation)
    3E6-3E7
    (determined by in situ hybridisation)
    3F1-3F5
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (47)
    Genomic Clones (26)
    cDNA Clones (119)
     

    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
    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
      Antibody Information
      Laboratory Generated Antibodies
       

      polyclonal

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

      Source for identity of: Tlk tlk

      Source for database merge of

      Source for merge of: tlk anon-WO03040301.114 anon-WO03040301.116

      Source for merge of: tlk CG32781

      Source for merge of: CG12462 EP1413

      Source for merge of: CG32782 CG33219

      Source for merge of: tlk CG12462

      Source for merge of: tlk CG32782

      Source for merge of: Tlk EP1408

      Additional comments

      Annotations CG32782 and CG32781 merged as CG34412 in release 5.2 of the genome annotation.

      Sequences corresponding to annotation CG12462 (which was eliminated in release 5.1 of the genome annotation) have been reinstated as part of the CG34412 annotation (which corresponds to tlk) in release 5.2 of the genome annotation.

      "l(1)G0299" may affect "CG2829". "l(1)G0167" may affect "CG2829". "l(1)G0343" may affect "CG2829". "l(1)G0054" may affect "CG2829".

      Source for merge of tlk anon-WO03040301.114 anon-WO03040301.116 was sequence comparison ( date:051113 ).

      Source for merge of tlk CG32782 was sequence comparison ( date:050324 ).

      Other Comments

      RNAi screen using dsRNA made from templates generated with primers directed against this gene causes decreased γ-tubulin staining at the spindle pole and misalignment of chromosomes along the spindle when assayed in S2 cells. This phenotype can be observed when the screen is performed with or without Cdc27 dsRNA.

      Annotation CG12462 eliminated in release 5.1 of the genome annotation.

      RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a greater than three-fold increase in AttA activity in response to heat-killed E.coli after ecdysone treatment in S2 cells.

      When dsRNA constructs are made and transiently transfected into S2 cells in RNAi experiments, an increase in the proportion of S phase cells, G2/M+, an increase in the proportion of G2/M phase cells, a decrease in cytokinetic index and a decrease in the mitotic index are seen.

      New annotation (CG33219) in release 3.2 of the genome annotation.

      The release 4 CG32782 annotation contains sequences corresponding to release 2 annotation CG2829. The release 3 version of CG32782 was incomplete and was missing most of the sequence corresponding to release 2 annotation CG2829 (possibly due to an assembly error), but this has been fixed in release 4 and the release 4 CG32782 annotation now includes all of the release 2 CG2829 sequence. 'CG33219' does not appear in Release 4. The region it was located in in Release 3.2 is now a large TE. 'CG33219' is homologous to annotation CG32782; presumably that is where this little bit of mislaid 'CG33219' sequence originated.

      The tlk-asf1 pathway is important for both DNA replication and chromatin assembly. tlk may coordinate cell cycle progression through the regulation of chromatin dynamics.

      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: cells become round and detached.

      New annotation (CG32782) in release 3 of the genome annotation. Annotation CG2829 was eliminated in release 3 of the genome annotation, October 2002.

      New annotation (CG32781) in release 3 of the genome annotation.

      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 73 )
      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.
      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
      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
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      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
      KEGG Genes - Molecular building blocks of life in the genomic space.
      MARRVEL_MODEL - MARRVEL (model organism gene)
      Linkouts
      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (23)
      Reported As
      Symbol Synonym
      BEST:GH17330
      BcDNA:GH07910
      EP1408
      anon-WO03040301.114
      anon-WO03040301.116
      Secondary FlyBase IDs
      • FBgn0086899
      • FBgn0052781
      • FBan0032782
      • FBgn0085441
      • FBgn0053219
      • FBgn0052782
      • FBgn0066856
      • FBgn0066855
      • FBgn0028489
      • FBgn0026698
      • FBgn0086709
      • FBgn0062688
      • FBgn0029675
      • FBgn0043460
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (118)