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General Information
Symbol
Dmel\dap
Species
D. melanogaster
Name
dacapo
Annotation Symbol
CG1772
Feature Type
FlyBase ID
FBgn0010316
Gene Model Status
Stock Availability
Gene Snapshot
dacapo (dap) encodes a Cyclin-dependent kinase inhibitor in the CIP/KIP family. It binds to CycE-Cdk2 complexes and thereby inhibits their protein kinase activity. Upregulation of dap is required after the last mitosis for arresting cells in G1/G0 before terminal differentiation in many post-mitotic cell types. [Date last reviewed: 2019-03-07]
Also Known As
Decapo, Cdi4, p21, chakra, p27
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:9,712,321..9,716,256 [+]
Recombination map
2-61
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
-
Summaries
Summary (Interactive Fly)
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\dap or the JBrowse view of Dmel\dap 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 5.42
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.49
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0088469
2530
245
Additional Transcript Data and Comments
Reported size (kB)
2.5 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0087553
27.1
245
10.27
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)
245 (aa); 27 (kD predicted)
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\dap using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (18 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (14 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from high throughput direct assay
inferred from high throughput direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
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
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

nurse cell | subset
nucleus

Comment: endocycling nurse cells

Additional Descriptive Data
dap transcript levels are at a constant low level in endocycling nurse cells.
Maternally derived dap transcripts are observed in early syncytial cycles then disappear rapidly in the syncytial blastoderm stage except in the pole cells. Zygotic expression begins during cellularization and persists during gastrulation around the amnioserosa. During germ band extension, expression is seen in the anterior and posterior midgut anlagen and in the developing CNS. In neuroblast lineages, dap expression appears to start just before the terminal division which generates neurons which exit from the mitotic cycle. In the epidermis, expression is at background levels and then expands greatly just before the onset of mitosis 16. Transcripts persist for about 2 hours after mitosis 16. dap transcripts are also observed in mesodermal cells that become postmitotic during these stages. After germ band retraction, expression is restricted mainly to the developing CNS and PNS. In summary, transient up-regulation of dap is tightly correlated with the arrest of cell proliferation in embryos. In larval eye imaginal discs, high levels of dap transcripts are observed in postmitotic cells of the differentiating ommatidial clusters posterior to the morphogenetic furrow.
dap transcripts are detected at the posterior pole of blastoderm embryos where the pole cells will form. At the onset of gastrulation, they are observed in the amnioserosa and in the cephalic and ventral furrows. After germ band elongation, they are expressed in dorsal and ventral cells of the epidermis. After germ band retraction, they are found in the CNS and PNS. After dorsal closure, dap is expressed in a small subset of cells in the ventral nerve cord and in the brain. In each of these cases, expression coincides with cell cycle arrest during development. Later in development, dap transcripts are detected in eye imaginal discs in the morphogenetic furrow and weakly posterior to the furrow.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
epidermis | restricted

Comment: S-phase of embryonic cycle 16

photoreceptor cell | subset of eye disc posterior to the morphogenetic furrow

Comment: photoreceptor precursors immediately posterior to morphogenetic furrow only

Additional Descriptive Data
In ovaries dap protein is first detected germarium region 2a. In maturing egg chambers, dap protein levels oscillate in endocycling nurse cells. Expression levels in the oocyte nucleus are constant.
The pattern of dap protein expression is similar to that of dap transcription.
dap protein expression correlates with exit of cells from the cell cycle. It is first observed in early embryos in yolk nuclei and later in the pole cells. During gastrulation, dap protein is expressed at high levels in the amnioserosa and is also seen in a row of cells on either side of the ventral furrow. After the onset of germ band elongation, dap expression is observed in the epidermis, first dorsally and then in dorsolateral cells and the ventral epidermis. During germ band retraction dap protein is expressed in cells of the PNS and CNS. In all cases, the peak of dap expression occurs as cells are completing their final divisions. In most cases, expression is maintained briefly and then subsides.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from high throughput direct assay
inferred from high throughput direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\dap 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
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
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, and Transgenic Constructs
Classical and Insertion Alleles ( 27 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 44 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of dap
Transgenic constructs containing regulatory region of dap
Deletions and Duplications ( 11 )
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
embryonic abdomen & glial cell | ectopic
external sensory organ precursor cell IIIb & S phase
follicle cell & nucleus | somatic clone, with Scer\GAL4Act5C.PP
G1 phase & external sensory organ precursor cell IIa, with Scer\GAL4neur-P72
G1 phase & external sensory organ precursor cell IIb, with Scer\GAL4neur-P72
neuron & embryonic thorax, with Scer\GAL4sca-537.4
PNS glial cell & S phase
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
Yes
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
Yes
Rattus norvegicus (Norway rat) (1)
1 of 13
Yes
Yes
Xenopus tropicalis (Western clawed frog) (1)
1 of 12
Yes
Yes
Danio rerio (Zebrafish) (3)
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (2)
2 of 15
Yes
Yes
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190F3Q )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
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) ( EOG09150BS4 )
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) ( EOG090W0NT6 )
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
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
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0O2O )
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
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (0)
No records found.
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 ( 3 )
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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
    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
    suppressible
    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
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-61
    Cytogenetic map
    Sequence location
    2R:9,712,321..9,716,256 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    46B1-46B1
    Limits computationally determined from genome sequence between P{lacW}l(2)k09221k09221&P{PZ}dap04454 and P{lacW}l(2)k05420k05420
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    46B1-46B2
    (determined by in situ hybridisation)
    46B-46B
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (12)
    Genomic Clones (14)
     

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

    cDNA Clones (85)
     

    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 sequences
    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
       
      Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
      Other Information
      Relationship to Other Genes
      Source for database identify of
      Source for database merge of
      Source for merge of: dap chakra
      Additional comments
      Other Comments
      dap is necessary for the maintenance of the meiotic cycle and differentiation state of oocytes.
      dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
      dap expression is controlled by modular arrays of tissue specific cis-regulatory elements.
      dpp does not mediate G1 arrest in the eye disc by inducing dap expression.
      Isolated by interaction with cdc2 and cdc2c using the interaction trap two hybrid system.
      dap is required to arrest cell proliferation in the embryonic epidermis at the correct developmental stage.
      Identified in a screen for genes that interact with R. dap is essential for normal embryonic development, it functions during embryogenesis to achieve a precisely timed exit from the cell cycle. Overexpression during eye development interferes with cell cycle progression and interacts genetically with Rbf and CycE. In absence of gross abnormalities in embryonic morphology or in cell fate determination the reason for lethality in dap embryos is unclear and may reflect additional functions of dap.
      dap is essential in controlling S phase entry.
      Yeast two-hybrid system has been used to characterise Cyclin-dependent kinase interactor proteins (the prey) in hunts with either cdc2 or cdc2c as baits. dap interacts with cdc2c and Cdi11, and other human cyclin-dependent kinases to activate transcription of a reporter gene, Ecol\lacZ.
      The yeast interaction trap selection has been used to isolate cell cycle regulatory proteins that interact with cdc2 and cdc2c. dap interacts preferentially with cdc2c.
      Origin and Etymology
      Discoverer
      Etymology
      "chakra" means "wheel" in Sanskrit.
      Identification
      External Crossreferences and Linkouts ( 52 )
      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/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
      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
      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
      Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
      FLIGHT - Cell culture data for RNAi and other high-throughput technologies
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      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 (30)
      Reported As
      Symbol Synonym
      CDKN2B
      E(Sev-CycE)2B
      cdi4
      dacapo/cyclin-dependent kinase interactor 4
      dap
      (Otsuki and Brand, 2019, Rodriguez-Fernandez et al., 2019, Thor, 2019.7.19, Akishina et al., 2018, Ariss et al., 2018, Mora et al., 2018, Spéder and Brand, 2018, Akishina et al., 2017, Cheng et al., 2017, Jin et al., 2017, Neto et al., 2017, Transgenic RNAi Project members, 2017-, Vazquez-Pianzola et al., 2017, Bivik et al., 2016, Shaikh et al., 2016, Aparicio et al., 2015, Barrios et al., 2015, Bivik et al., 2015, Bouleau and Tricoire, 2015, Dequéant et al., 2015, Liu et al., 2015, Reimels and Pfleger, 2015, Sanchez-Alvarez et al., 2015, Sun and Buttitta, 2015, Venkei and Yamashita, 2015, Wang and Baker, 2015, Ciglar et al., 2014, Gómez-Lamarca et al., 2014, Wong et al., 2014, Iovino et al., 2013, Lim et al., 2013, Manansala et al., 2013, Schertel et al., 2013, Zhai et al., 2012, Colonques et al., 2011, Ghorbani et al., 2011, Guest et al., 2011, Aerts et al., 2010, Baig et al., 2010, Fan et al., 2010, Herranz et al., 2010, Kong et al., 2010, Reddy et al., 2010, zur Lage and Jarman, 2010, Bossuyt et al., 2009, Dubruille et al., 2009, Narbonne-Reveau and Lilly, 2009, Nicholson et al., 2009, Read et al., 2009, Rogers et al., 2009, Southall and Brand, 2009, Wang and Kalderon, 2009, Yu et al., 2009, Caussinus et al., 2008, Gupta et al., 2008, Remaud et al., 2008, Sukhanova and Du, 2008, Buszczak et al., 2007, Buttitta et al., 2007, Escudero and Freeman, 2007, Garcia et al., 2007, Griffiths et al., 2007, Hong et al., 2007, Junion et al., 2007, Kugler and Nagel, 2007, Nurminsky, 2007, Quinones-Coello, 2007, Sandmann et al., 2007, Secombe et al., 2007, Shibutani et al., 2007, Steinhilb et al., 2007, Sukhanova et al., 2007, Tyler et al., 2007, Khurana et al., 2006, Legent et al., 2006, Morris et al., 2006, Mukherjee et al., 2006, Parker, 2006, Reber et al., 2006, Firth and Baker, 2005, Brodsky et al., 2004, Delanoue et al., 2004, Heidmann et al., 2004, Stanyon et al., 2004, Heriche et al., 2003, Huet et al., 2002, Gim et al., 2001, Wallace et al., 2000)
      l(2)04454
      p21dacapo
      Name Synonyms
      Cyclin-dependent kinase interactor 4
      Dacapo/p21
      p21/dacapo
      Secondary FlyBase IDs
      • FBgn0010576
      • FBgn0015920
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (323)