FB2025_04 , released October 2, 2025
Gene: Dmel\dap
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General Information
Symbol
Dmel\dap
Species
D. melanogaster
Name
dacapo
Annotation Symbol
CG1772
Feature Type
protein_coding_gene
FlyBase ID
FBgn0010316
Gene Model Status
Current
Stock Availability
37 publicly available
Gene Summary
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] (FlyBase Gene Snapshot)
All Summaries
Also Known As

Decapo, p21, Cdi4, p27Dap, chakra

Key Links
Genomic Location
Cytogenetic map
46B1-46B1
Sequence location
Recombination map
2-61
RefSeq locus
NT_033778 REGION:9712321..9716256
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (15 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables cyclin-dependent protein serine/threonine kinase inhibitor activity
inferred from mutant phenotype
(Narbonne-Reveau and Lilly, 2009)
inferred from direct assay
(de Nooij et al., 1996, Lane et al., 1996)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables cyclin-dependent protein serine/threonine kinase inhibitor activity
inferred from electronic annotation with InterPro:IPR003175
(InterPro Project Members, 2004-)
Biological Process (12 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
involved_in germarium-derived oocyte fate determination
inferred from mutant phenotype
(Hong et al., 2003)
involved_in germline cell cycle switching, mitotic to meiotic cell cycle
inferred from mutant phenotype
(Narbonne-Reveau and Lilly, 2009)
involved_in histoblast morphogenesis
inferred from mutant phenotype
(Ninov et al., 2007)
involved_in mitotic cell cycle
inferred from mutant phenotype
(Parker, 2006)
involved_in negative regulation of G1/S transition of mitotic cell cycle
inferred from mutant phenotype
(Firth and Baker, 2005)
involved_in negative regulation of mitotic cell cycle
inferred from direct assay
(de Nooij et al., 1996, Lane et al., 1996)
involved_in premeiotic DNA replication
inferred from mutant phenotype
(Narbonne-Reveau and Lilly, 2009)
involved_in sensory organ precursor cell division
inferred from mutant phenotype
(Audibert et al., 2005)
involved_in ventral cord development
inferred from high throughput mutant phenotype
(Bivik et al., 2015)
involved_in wing disc development
inferred from genetic interaction with FLYBASE:ab; FB:FBgn0264442
(Simoes da Silva et al., 2019)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
involved_in negative regulation of mitotic cell cycle
inferred from biological aspect of ancestor with PANTHER:PTN004142838
(GO Reference Genome Project, 2011-)
involved_in oogenesis
traceable author statement
(Lee and Orr-Weaver, 2003)
involved_in regulation of cell cycle
inferred from electronic annotation with InterPro:IPR003175
(InterPro Project Members, 2004-)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in nucleoplasm
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleus
inferred from direct assay
(Lane et al., 1996)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
located_in nucleus
inferred from electronic annotation with InterPro:IPR003175
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
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]
Summary (Interactive Fly)

CIP/KIP family cyclin dependent kinase inhibitor - inhibits cell cycle progression - .binds to CycE-Cdk2 complexes - Upregulation of Dap is required after the last mitosis for arresting cells in G1/G0 before terminal differentiation in many post-mitotic cell types

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

Please see 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
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q7JNL9)

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

Gene model reviewed during 5.42

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

Gene model reviewed during 5.49

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
dap-RA
FBtr0088469
NM_057600
2530
245
Additional Transcript Data and Comments
Reported size (kB)

2.5 (northern blot)

(de Nooij et al., 1996)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
dap-PA
FBpp0087553
27.1
245
10.27
Q7JNL9
NP_476948
AAF58907
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)

(Lane et al., 1996)

245 (aa); 27 (kD predicted)

(de Nooij et al., 1996)
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
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).

-0.32

Transcript Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
oogenesis & adult stage | female
oocyte
• nucleus
(de Nooij et al., 2000)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
anterior midgut primordium
(Lane et al., 1996)
larval ventral nerve cord | restricted
(de Nooij et al., 1996)
presumptive embryonic/larval central nervous system
(de Nooij et al., 1996, Lane et al., 1996)
epidermis
(de Nooij et al., 1996)
ectoderm
(Lane et al., 1996)
supraesophageal ganglion
(de Nooij et al., 1996)
mesoderm
(Lane et al., 1996)
posterior midgut primordium
(Lane et al., 1996)
peripheral nervous system
(de Nooij et al., 1996, Lane et al., 1996)
amnioserosa
(Lane et al., 1996)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 2
organism
(Lane et al., 1996)
blastoderm stage
organism | 0-10% egg length
(de Nooij et al., 1996)
embryonic stage 4 -- 6
anterior endoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
endoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
primordial germ cell
(Fisher et al., 2012)
pole plasm
(Fisher et al., 2012)
embryonic stage 6
ventral furrow
(de Nooij et al., 1996)
amnioserosa
(de Nooij et al., 1996)
cephalic furrow
(de Nooij et al., 1996)
embryonic stage 7
amnioserosa
(Liu et al., 2002)
embryonic stage 7 -- 8
amnioserosa anlage
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic stage 9 -- 10
amnioserosa primordium
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic cycle 16
presumptive embryonic/larval central nervous system | restricted

Comment: S-phase

(de Nooij et al., 2000)
presumptive embryonic/larval peripheral nervous system | restricted

Comment: S-phase

(de Nooij et al., 2000)
embryonic epidermis | restricted

Comment: S-phase

(de Nooij et al., 2000)
embryonic stage 11
tracheal pit

Comment: early stage 11

(Liu et al., 2002)
embryonic epidermis

Comment: late stage 11

(Liu et al., 2002)
embryonic stage 11 -- 12
epidermis | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
amnioserosa
(Fisher et al., 2012)
anterior midgut primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
tracheal primordium
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
embryonic stage 13
presumptive embryonic/larval peripheral nervous system
(Liu et al., 2002)
embryonic stage 13 -- 14
presumptive embryonic/larval central nervous system | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
presumptive embryonic/larval peripheral nervous system | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
embryonic stage 13 -- 16
presumptive embryonic/larval central nervous system
(Liu et al., 2002)
dorsal sensory complex proper primordium

Comment: reported as dorsal/lateral sensory complexes

(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic head sensory system
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
ventral sensory complex primordium
(Fisher et al., 2012)
larval stage
eye disc morphogenetic furrow
(de Nooij et al., 1996)
eye-antennal disc
(de Nooij et al., 1996, Lane et al., 1996)
eye disc posterior to the morphogenetic furrow
(Lane et al., 1996)
third instar larval stage
larval inner optic anlage
(Liu et al., 2002, Wallace et al., 2000)
larval outer optic anlage
(Liu et al., 2002)
lamina furrow
(Liu et al., 2002)
eye disc morphogenetic furrow
(Baig et al., 2010, Liu et al., 2002)
photoreceptor neuron | subset of eye disc posterior to the morphogenetic furrow

Comment: photoreceptor precursors immediately posterior to morphogenetic furrow only

(de Nooij et al., 2000)
lamina anlage | variable
(Wallace et al., 2000)
adult stage | female & oogenesis
nurse cell | subset
• nucleus

Comment: endocycling nurse cells

(de Nooij et al., 2000)
Additional Descriptive Data

dap transcript levels are at a constant low level in endocycling nurse cells.

(de Nooij et al., 2000)

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.

(Lane et al., 1996)

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.

(de Nooij et al., 1996)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
larval ventral nerve cord | restricted
(de Nooij et al., 1996)
epidermis
(de Nooij et al., 1996)
presumptive embryonic/larval central nervous system
(de Nooij et al., 1996)
peripheral nervous system
(de Nooij et al., 1996)
embryonic stage 4
primordial germ cell
(de Nooij et al., 1996)
yolk nucleus
(de Nooij et al., 1996)
embryonic stage 6
amnioserosa
(de Nooij et al., 1996)
embryonic cycle 16
presumptive embryonic/larval central nervous system | restricted

Comment: S-phase

(de Nooij et al., 2000)
presumptive embryonic/larval peripheral nervous system | restricted

Comment: S-phase

(de Nooij et al., 2000)
embryonic epidermis | restricted

Comment: S-phase

(de Nooij et al., 2000)
embryonic stage 11 -- 12
epidermis | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
embryonic stage 13 -- 14
presumptive embryonic/larval central nervous system | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
presumptive embryonic/larval peripheral nervous system | restricted

Comment: S-phase of embryonic cycle 16

(de Nooij et al., 2000)
third instar larval stage
photoreceptor neuron | subset of eye disc posterior to the morphogenetic furrow

Comment: photoreceptor precursors immediately posterior to morphogenetic furrow only

(de Nooij et al., 2000)
adult stage | female & oogenesis
germarium region 2a
(de Nooij et al., 2000)
nurse cell | subset
• nucleus

Comment: endocycling nurse cells

(de Nooij et al., 2000)
oogenesis & adult stage | female
oocyte
• nucleus
(de Nooij et al., 2000)
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.

(de Nooij et al., 2000)

The pattern of dap protein expression is similar to that of dap transcription.

(Lane et al., 1996)

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.

(de Nooij et al., 1996)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in nucleoplasm
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleus
inferred from direct assay
(Lane et al., 1996)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\dap in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
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
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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 29 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 46 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of dap
Transgenic constructs containing regulatory region of dap
Aberrations (Deficiencies and Duplications) ( 11 )
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
lethal
lethal, with Scer\GAL4ap-md544
lethal, with Scer\GAL4Dll-md23
lethal, with Scer\GAL4T80
lethal, with Scer\GAL4αTub84B.PL
lethal (with Df(2R)BSC131)
lethal (with Df(2R)BSC132)
lethal | recessive
partially lethal
partially lethal - majority die
partially lethal - majority die, with Scer\GAL432B
viable
Sterility
Allele
female sterile, with Scer\GAL4c323
Other Phenotypes
Allele
abnormal cell cycle, with Scer\GAL4en.PU
abnormal cell cycle, with Scer\GAL4hh.PU
abnormal endomitotic cell cycle | adult stage | female, with Scer\GAL4c323
abnormal endomitotic cell cycle | embryonic stage, with Scer\GAL4αTub84B.PL
abnormal endomitotic cell cycle | third instar larval stage, with Scer\GAL4ptc-559.1
abnormal mitotic cell cycle
abnormal mitotic cell cycle, with Scer\GAL4en.PU
abnormal mitotic cell cycle, with Scer\GAL4neur-P72
abnormal mitotic cell cycle | embryonic stage, with Scer\GAL4en-e16E
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4sca-537.4
abnormal neuroanatomy | embryonic stage
abnormal neuroanatomy | embryonic stage | recessive
abnormal size, with Scer\GAL4ey.PH
abnormal size | adult stage, with Scer\GAL4ey.PH
abnormal size | third instar larval stage, with Scer\GAL4ey.PH
decreased cell death | embryonic stage, with Scer\GAL4en-e16E
decreased cell number | adult stage | conditional, with Scer\GAL4byn-Gal4
decreased occurrence of cell division
decreased occurrence of cell division | larval stage, with Scer\GAL4en-e16E
decreased occurrence of cell division | larval stage, with Scer\GAL4GMR.PU
decreased occurrence of cell division | larval stage, with Scer\GAL4ptc-559.1
increased cell number | embryonic stage
increased cell number | embryonic stage | recessive
increased occurrence of cell division | embryonic stage
increased occurrence of cell division | larval stage
increased occurrence of cell division | larval stage, with Scer\GAL4wor.PA
visible
visible, with Scer\GAL4en-e16E
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4GMR.PF
visible, with Scer\GAL4ptc-559.1
visible, with Scer\GAL4salm.EPv
visible, with Scer\GAL4sca-537.4
visible, with Scer\GAL4sd.PU
visible, with Scer\GAL4vg.int2.1
visible, with Scer\GAL4vg.PU
visible | adult stage, with Scer\GAL4en-e16E
visible | adult stage, with Scer\GAL4nub-AC-62
Phenotype manifest in
Allele
adult abdomen | somatic clone, with Scer\GAL4Act5C.PI
adult pyloric epithelium | conditional, with Scer\GAL4byn-Gal4
apterous-expressing neuron of the lateral cluster of the thorax | ectopic
apterous-expressing neuron of the lateral cluster of the thorax Ap4 | increased number
chaeta, with Scer\GAL4pnr-MD237
dorsal appendage, with Scer\GAL4c323
dorsal vessel wall cell, with Scer\GAL4twi.PG
dorsocentral bristle, with Scer\GAL4sca-537.4
egg, with Scer\GAL4c323
egg chorion, with Scer\GAL4c323
embryo
embryonic/larval epidermis | somatic clone, with Scer\GAL4Act5C.PI
embryonic/larval midgut | embryonic stage, with Scer\GAL4αTub84B.PL
embryonic/larval optic lobe | third instar larval stage
embryonic/larval salivary gland | third instar larval stage, with Scer\GAL4ptc-559.1
embryonic/larval tracheal dorsal trunk
embryonic/larval tracheal transverse connective
embryonic abdomen & glial cell | ectopic
embryonic epidermis
embryonic longitudinal glial cell
embryonic pericardial cell, with Scer\GAL4eve.eme
embryonic pericardial cell, with Scer\GAL4twi.PG
embryonic segment posterior compartment, with Scer\GAL4en-e16E
external sensory organ precursor cell IIIb & S phase
eye
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4GMR.PF
eye disc, with Scer\GAL4ey.PH
eye disc posterior to the morphogenetic furrow | somatic clone, with Scer\GAL4GMR.PU
follicle cell & nucleus | somatic clone, with Scer\GAL4Act5C.PP
follicle stem cell | somatic clone
G1 phase & external sensory organ precursor cell IIa, with Scer\GAL4neur-P72
G1 phase & external sensory organ precursor cell IIb, with Scer\GAL4neur-P72
histoblast | somatic clone, with Scer\GAL4Act5C.PI
karyosome | germline clone
larval central brain, with Scer\GAL4wor.PA
larval EL neuron | embryonic stage | increased number
larval neuroblast, with Scer\GAL4wor.PA
larval neuroblast | embryonic stage
lch1 neuron
macrochaeta
macrochaeta | ectopic
main body follicle cell, with Scer\GAL4c323
mesonotum, with Scer\GAL4pnr-MD237
mitotic cell cycle, with Scer\GAL4Act5C.PP
mitotic cell cycle, with Scer\GAL4unspecified
mitotic cell cycle | ectopic
neuron & embryonic thorax, with Scer\GAL4sca-537.4
nucleus | third instar larval stage, with Scer\GAL4ptc-559.1
ommatidium
oocyte | germline clone
PNS glial cell & S phase
scutellar bristle, with Scer\GAL4sca-537.4
thoracic neuroblast NB6-4, with Scer\GAL4sca-537.4
tormogen cell, with Scer\GAL4sca-537.4
trichogen cell, with Scer\GAL4sca-537.4
ventral nerve cord primordium | embryonic stage
wing, with Scer\GAL4salm.EPv
wing, with Scer\GAL4sd.PU
wing, with Scer\GAL4vg.int2.1
wing blade posterior compartment, with Scer\GAL4en-e16E
wing cell, with Scer\GAL4en-e16E
wing disc, with Scer\GAL4ptc-559.1
wing hair, with Scer\GAL4en-e16E
wing hair, with Scer\GAL4nub-AC-62
wing margin, with Scer\GAL4nub-AC-62
wing margin, with Scer\GAL4ptc-559.1
wing margin, with Scer\GAL4vg.PU
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (3)
Hsap\CDKN1B
4 of 14
Yes
Yes
3  
Hsap\CDKN1C
4 of 14
Yes
Yes
6  
Hsap\CDKN1A
2 of 14
No
Yes
7  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (3)
Rnor\Cdkn1b
4 of 14
Yes
Yes
Rnor\Cdkn1a
2 of 14
No
Yes
Rnor\Cdkn1c
2 of 14
No
Yes
Mus musculus (laboratory mouse) (3)
Mmus\Cdkn1b
3 of 14
Yes
Yes
Mmus\Cdkn1c
3 of 14
Yes
Yes
Mmus\Cdkn1a
2 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
Xtro\cdkn1a
1 of 13
Yes
No
Xtro\cdkn1c
1 of 13
Yes
Yes
Xtro\cdknx
1 of 13
Yes
Yes
Danio rerio (Zebrafish) (6)
Drer\cdkn1ba
4 of 14
Yes
Yes
Drer\cdkn1bb
4 of 14
Yes
Yes
Drer\cdkn1ca
3 of 14
No
Yes
Drer\cdkn1a
2 of 14
No
Yes
Drer\cdkn1cb
1 of 14
No
Yes
Drer\cdkn1d
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (2)
Cele\cki-1
6 of 14
Yes
Yes
Cele\cki-2
4 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP010629
1 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:dap. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
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 ( 3 )
    Modifiers Based on Experimental Evidence ( 6 )
    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?
    CDKN1B; cyclin dependent kinase inhibitor 1B
    4 of 14
    600778
     
    CDKN1C; cyclin dependent kinase inhibitor 1C
    4 of 14
    CYCLIN-DEPENDENT KINASE INHIBITOR 1C; CDKN1C
     
    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
    Interaction Browsers
    View in FlyBase Interactions Browser   View in MIST tool (external link)
    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    dap
    enhanceable
    CycE
    (Staehling-Hampton et al., 1999)
    dap
    suppressible
    CycE
    (Audibert et al., 2005, Lane et al., 1996)
    dap|Rbf
    suppressible
    E2f1
    (Weng et al., 2003)
    Rbf|dap
    suppressible
    E2f1
    (Weng et al., 2003)
    dap|CycE
    suppressible
    spen
    (Staehling-Hampton et al., 1999)
    dap
    enhanceable
    vg
    (Legent et al., 2006)
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    aurA
    suppressible
    dap
    (Wang et al., 2006)
    bsk|BubR1
    suppressible
    dap
    (Wong et al., 2014)
    caup
    enhanceable
    dap
    (Barrios et al., 2015)
    ci
    suppressible
    dap
    (Duman-Scheel et al., 2002)
    CycE
    enhanceable
    dap
    (Wang and Kalderon, 2009, Lane et al., 2000, Secombe et al., 1998)
    CycE
    suppressible
    dap
    (Wang and Kalderon, 2009, Audibert et al., 2005, Hong et al., 2003, Secombe et al., 1998)
    Dcr-1
    suppressible
    dap
    (Herranz et al., 2010, Hatfield et al., 2005)
    Delta|lola|psq
    suppressible
    dap
    (Bossuyt et al., 2009)
    H
    suppressible
    dap
    (Müller et al., 2005)
    InR
    suppressible
    dap
    (Yu et al., 2009)
    kuz
    suppressible
    dap
    (Bivik et al., 2016)
    N
    enhanceable
    dap
    (Sood et al., 2022)
    p53
    suppressible
    dap
    (Fan et al., 2010)
    Pi3K92E|btl::Egfr
    suppressible
    dap
    (Read et al., 2009)
    png
    suppressible
    dap
    (Lee et al., 2001)
    pzg
    suppressible
    dap
    (Kugler and Nagel, 2007)
    Rap1
    enhanceable
    dap
    (de Nooij et al., 1996)
    Ras85D
    enhanceable
    dap
    (Ito and Igaki, 2021)
    Ras85D|scrib
    suppressible
    dap
    (Brumby and Richardson, 2003)
    Rbf
    enhanceable
    dap
    (Weng et al., 2003, Xin et al., 2002)
    Rbf
    suppressible
    dap
    (Xin et al., 2002)
    Rheb
    suppressible
    Rbf|dap
    (Khurana et al., 2006)
    Skp2
    suppressible
    dap
    (Dui et al., 2013)
    upd1
    suppressible
    dap
    (Mukherjee et al., 2006)
    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)
    2R
    Recombination map
    2-61
    Cytogenetic map
    46B1-46B1
    Sequence location
    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)
    (Abdelilah-Seyfried et al., 2000, Spradling et al., 1999)
    46B1-46B14
    (de Nooij, 1996.11.12)
    46B-46B
    (determined by in situ hybridisation)
    (de Nooij et al., 1996, Lane et al., 1996)
    46B1-46B2
    (BDGP Project Members, 1994-1999)
    Experimentally Determined Recombination Data
    Location

    2-61

    (FlyBase, 2016)

    2-59.7

    (Comeron et al., 2012)

    2-

    (de Nooij, 1996.11.12)
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (37)
    Genomic Clones (14)
     

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

    cDNA Clones (82)
     

    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)
      BDGP DGC clones
      Other clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
       

      polyclonal

      (Baumgardt et al., 2014, de Nooij et al., 1996)
      Commercially Available Antibodies
       
      Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          dap is necessary for the maintenance of the meiotic cycle and differentiation state of oocytes.

          (Hong et al., 2003)

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

          (Kiger et al., 2003)

          dap expression is controlled by modular arrays of tissue specific cis-regulatory elements.

          (Liu et al., 2002)

          dpp does not mediate G1 arrest in the eye disc by inducing dap expression.

          (Horsfield et al., 1998)

          Isolated by interaction with cdc2 and cdc2c using the interaction trap two hybrid system.

          (Finley et al., 1996)

          dap is required to arrest cell proliferation in the embryonic epidermis at the correct developmental stage.

          (Lane et al., 1996)

          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.

          (de Nooij et al., 1996)

          dap is essential in controlling S phase entry.

          (de Nooij, 1996.4.29)

          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.

          (Finley and Brent, 1994)

          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.

          (Finley and Brent, 1993)
          Relationship to Other Genes
          Source for database merge of

          Source for merge of: dap chakra

          (Hariharan, 2005.5.18)
          Additional comments
          Nomenclature History
          Source for database identify of
          Nomenclature comments
          Etymology

          "chakra" means "wheel" in Sanskrit.

          (de Nooij, 1996.4.29)
          Synonyms and Secondary IDs (30)
          Reported As
          Symbol Synonym
          CDI4
          (Finley and Brent, 1993)
          CDKN2B
          (Bier, 2005)
          CG1772
          (Meadows, 2019.11.20, Schertel et al., 2013, Ni et al., 2011.7.19, Aerts et al., 2010, Bossuyt et al., 2009, Lebo et al., 2009, Perkins et al., 2009, Quinones-Coello, 2007, Norga et al., 2003)
          CIB1
          (Bier, 2005)
          Cdi4
          (Finley et al., 1996, Finley and Brent, 1994)
          DAP
          (Al Zouabi et al., 2023, Mirkovic et al., 2019, Mbodj et al., 2013)
          Dac
          (Higa et al., 2006)
          Dacapo
          (Clemente and Weavers, 2023, Neto et al., 2017)
          Dap
          (Almeida Machado Costa et al., 2022, Ray and Li, 2022, Ray et al., 2022, Doherty et al., 2021, Kim et al., 2021, Ding et al., 2020, Maurange, 2020, Bivik Stadler et al., 2019, Curt et al., 2019, Toshniwal et al., 2019, Yadav et al., 2016, Li et al., 2015, Swanson et al., 2015, Zielke et al., 2014, Dui et al., 2013, Gordon et al., 2013, Guest et al., 2011, Ambrus and Frolov, 2010, Narbonne-Reveau and Lilly, 2009, Read et al., 2009, Bowman et al., 2008, Ninov et al., 2007, Steinhilb et al., 2007, Sukhanova et al., 2007, Fichelson et al., 2005, Mandal et al., 2005, Reis and Edgar, 2003)
          Decapo
          (Miles et al., 2011, Bier, 2005)
          E(Sev-CycE)2B
          (Lane et al., 2000)
          P15
          (Bier, 2005)
          cdi4
          dacapo
          (Corrales et al., 2022, Morelli et al., 2018, Gogendeau et al., 2015)
          dacapo/cyclin-dependent kinase interactor 4
          (Li and White, 2003)
          dap
          (Datta and Bangi, 2024, Sood et al., 2024, Trujillo et al., 2024, Zhou et al., 2024, Chan et al., 2023, Titlow et al., 2023, Zhang et al., 2023, Diegmiller et al., 2022, Doherty et al., 2022, Bilder et al., 2021, Ferguson et al., 2021, Ito and Igaki, 2021, Joy et al., 2021, Khan et al., 2021, Michki et al., 2021, Neiswender et al., 2021, Tatapudy et al., 2021, Bahrampour and Thor, 2020, Hassan et al., 2020, Melamed and Kalderon, 2020, Morita et al., 2020, Port et al., 2020, Yaghmaeian Salmani and Thor, 2020, Harding and White, 2019, Otsuki and Brand, 2019, Rodriguez-Fernandez et al., 2019, Shcherbata, 2019, Simoes da Silva et al., 2019, Thor, 2019.7.19, van den Ameele and Brand, 2019, Yee et al., 2019, 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, Inaba 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, Nakajima 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
          p21
          (Khan et al., 2019, Meserve and Duronio, 2017, Gupta et al., 2008, Jordan et al., 2006)
          p21dacapo
          (Coffman, 2004)
          p27
          (Bouleau and Tricoire, 2015, Mandal et al., 2010)
          p27Dap
          (Shibutani et al., 2007, Swanhart et al., 2007)
          p27cip/kip
          (Southall et al., 2014)
          Name Synonyms
          Cyclin-dependent kinase interactor 4
          Dacap0
          (Escudero and Freeman, 2007)
          Dacapo
          (Bhattacharya et al., 2024, García-Puga et al., 2022, Soni et al., 2021, Bahrampour and Thor, 2020, Prat-Rojo et al., 2020, Neves and Eisenman, 2019, Banerjee and Roy, 2017, Martins et al., 2017, Kotov et al., 2016, Gupte, 2015, Dui et al., 2013, Manansala et al., 2013, Guest et al., 2011, Klusza and Deng, 2011, Ouyang et al., 2011, Ambrus and Frolov, 2010, Herranz et al., 2010, Monk et al., 2010, Reddy et al., 2010, Narbonne-Reveau and Lilly, 2009, Wang and Kalderon, 2009, Bowman et al., 2008, Owusu-Ansah et al., 2008, Remaud et al., 2008, Sukhanova and Du, 2008, Hong et al., 2007, Owusu-Ansah et al., 2007, Shibutani et al., 2007, Steinhilb et al., 2007, Sukhanova et al., 2007, Buttitta and Edgar, 2006, Jordan et al., 2006, Siddall et al., 2006, Wang et al., 2006, Baonza and Freeman, 2005, LeBrasseur, 2005, Delanoue et al., 2004, Gim et al., 2001)
          Dacapo/p21
          (Reynolds, 2007)
          Dacpao
          (Higa et al., 2006)
          Decapo
          (Chia et al., 2008, Khurana et al., 2006)
          chakra
          (de Nooij, 1996.4.29)
          dacapo
          (Morita et al., 2020, Otsuki and Brand, 2020, Spéder and Brand, 2018, Gupta et al., 2016, Aparicio et al., 2015, Barrios et al., 2015, Dekanty et al., 2015, Dequéant et al., 2015, Liu et al., 2015, Lucas et al., 2015, Powell et al., 2015, Reimels and Pfleger, 2015, Venkei and Yamashita, 2015, Gómez-Lamarca et al., 2014, Southall et al., 2014, Laver et al., 2013, Adamson and Lajeunesse, 2012, Guest et al., 2011, Buttitta et al., 2010, Fan et al., 2010, Mandal et al., 2010, zur Lage and Jarman, 2010, Bossuyt et al., 2009, Lebo et al., 2009, Rogers et al., 2009, Caussinus et al., 2008, Sato et al., 2008, Buttitta et al., 2007, Griffiths et al., 2007, Rani et al., 2007, Secombe et al., 2007, Tountas and Fortini, 2007, Brun et al., 2006, Hollien and Weissman, 2006, Legent et al., 2006, Morris et al., 2006, Mukherjee et al., 2006, Zielke et al., 2006, Brodsky et al., 2004, Gullaud et al., 2003, Delanoue et al., 2002, van de Bor et al., 1999, de Nooij, 1996.11.12, Finley, 1996.7.29)
          decapo
          (Slattery et al., 2011, Nicholson et al., 2009, Tower, 2004)
          p21/Dacapo
          (Reynolds et al., 2008)
          p21/dacapo
          (Yu et al., 2008)
          Secondary FlyBase IDs
          • FBgn0010576
          • FBgn0015920
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          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/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/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.
          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
          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
          Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
          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
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          References (389)