FB2026_02 , released June 18, 2026
Gene: Dmel\dpa
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General Information
Symbol
Dmel\dpa
Species
D. melanogaster
Name
disc proliferation abnormal
Annotation Symbol
CG1616
Feature Type
FlyBase ID
FBgn0015929
Gene Model Status
Stock Availability
Enzyme Name (EC)
DNA helicase (3.6.4.12)
DNA 3'-5' helicase (5.6.2.4)
Gene Summary
Acts as a component of the Mcm2-7 complex (Mcm complex) which is the putative replicative helicase essential for 'once per cell cycle' DNA replication initiation and elongation in eukaryotic cells. The active ATPase sites in the Mcm2-7 ring are formed through the interaction surfaces of two neighboring subunits such that a critical structure of a conserved arginine finger motif is provided in trans relative to the ATP-binding site of the Walker A box of the adjacent subunit. The six ATPase active sites, however, are likely to contribute differentially to the complex helicase activity. Required for DNA replication and cell proliferation. Essential role in mitotic DNA replication but not in endoreplication. (UniProt, Q26454)
Contribute a Gene Snapshot for this gene.
Also Known As

Mcm4, DmMCM4, l(2)43Ca

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-56
RefSeq locus
NT_033778 REGION:7496702..7500145
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (14 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
Biological Process (6 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
involved_in DNA replication
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR008047
involved_in DNA replication
inferred from electronic annotation with InterPro:IPR018525
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000179968
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
part_of CMG complex
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
part_of MCM complex
inferred from electronic annotation with InterPro:IPR008047
inferred from biological aspect of ancestor with PANTHER:PTN000179711
Protein Family (UniProt)
Belongs to the MCM family. (Q26454)
Catalytic Activity (EC/Rhea)
DNA helicase activity
ATP + H2O = ADP + phosphate + H(+) (3.6.4.12)
3'-5' DNA helicase activity
Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction (5.6.2.4)
Summaries
Gene Group (FlyBase)
MCM2-7 COMPLEX -
The heterohexameric MCM2-7 complex is the helicase motor of the CMG (Cdc45-MCM-GINS) replicative helicase of eukaryotes. The six MCM subunits are related to one another and are members of the AAA+ (ATPases associated with diverse cellular activities) superfamily. Hydrolysis of ATP by the individual MCM subunits catalyses the conformational changes that drive unwinding of the DNA duplex. However, the eukaryotic MCM complex alone has no or little helicase activity, requiring the other components of the CMG complex (Cdc45 and the GINS complex) for activation. (Adapted from PMID:23412083.)
CMG COMPLEX -
The CMG (Cdc45-MCM-GINS) complex is the eukaryotic replicative helicase, the enzyme that unwinds double-stranded DNA at replication forks. It is a macromolecular assembly of three replication factors: the Cdc45 protein, and the MCM and GINS complexes. All three components are essential for CMG function, with the MCM2-7 complex functioning as the molecular motor that harnesses the energy of ATP hydrolysis to catalyse strand separation. (Adapted from PMID:23412083.)
MCM FAMILY HELICASES -
The MCM (minichromosome maintenance) family comprise DNA helicases sharing a common 'MCM domain'. MCM2-7 form a heterohexameric complex that is essential for initiation of DNA replication and also acts during elongation as the replicative helicase. MCM8 and MCM9 are not part of this complex - they have roles in replication elongation, meiotic recombination and DNA repair. MCM8 and 9 arose early during eukaryotic evolution, though one or both have been lost in multiple lineages: in mammals, MCM8 and MCM9 form a functional heterodimeric complex; D. melanogaster and related species possess MCM8 only; while other species (e.g. yeast, C. elegans) have neither. (Adapted from PMID:23412083, PMID:30743181 and FBrf0188222.)
Protein Function (UniProtKB)
Acts as a component of the Mcm2-7 complex (Mcm complex) which is the putative replicative helicase essential for 'once per cell cycle' DNA replication initiation and elongation in eukaryotic cells. The active ATPase sites in the Mcm2-7 ring are formed through the interaction surfaces of two neighboring subunits such that a critical structure of a conserved arginine finger motif is provided in trans relative to the ATP-binding site of the Walker A box of the adjacent subunit. The six ATPase active sites, however, are likely to contribute differentially to the complex helicase activity. Required for DNA replication and cell proliferation. Essential role in mitotic DNA replication but not in endoreplication.
(UniProt, Q26454)
Summary (Interactive Fly)

Licensing factor - Zn finger motif - cdc21 homolog - MCM4 homolog - implicated in the initiation of DNA replication and in the discrimination between replicated and unreplicated chromatin

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\dpa 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 Q26454)

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
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Gene model reviewed during 5.48

Gene model reviewed during 5.56

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0088982
2803
866
FBtr0345797
3071
866
Additional Transcript Data and Comments
Reported size (kB)

2.8 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0088055
96.6
866
7.57
FBpp0311783
96.6
866
7.57
Polypeptides with Identical Sequences

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

866 aa isoforms: dpa-PA, dpa-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

866 (aa); 97 (kD predicted)

Comments
External Data
Subunit Structure (UniProtKB)

Component of the Mcm2-7 complex. The complex forms a toroidal hexameric ring with the proposed subunit order Mcm2-Mcm6-Mcm4-Mcm7-Mcm3-Mcm5 (Probable).

(UniProt, Q26454)
Post Translational Modification

Phosphorylated by the catalytic component of the Dbf4-dependent kinase (DDK) complex Cdc7.

(UniProt, Q26454)
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\dpa 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).

-0.04

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism | ubiquitous

Comment: maternally deposited

antennal primordium

Comment: reported as procephalic ectoderm primordium

central brain primordium

Comment: reported as procephalic ectoderm primordium

visual primordium

Comment: reported as procephalic ectoderm primordium

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

Additional Descriptive Data

dpa transcripts are detected in embryos from 0-12hrs with a peak in 6-9hr embryos and decline to very low levels thereafter. dpa is expressed in a pattern that closely resembles the pattern of DNA replication. There is expression in germline cells throughout oogenesis. The transcript accumulates in nurse cells late in oogenesis and then is transferred to the oocyte. The pattern of expression in embryos is very dynamic during gastrulation and germ band extension. Most tissues express the transcript at high levels but not all. Epidermal expression remains high until stage 11 at which point it is extinguished. Expression is then found mainly in the delaminated neuroblasts of the CNS and in the sensory organ precursors of the PNS while they are dividing. Expression in the brain continues until the end of embryogenesis. dpa is also expressed in endoreplicating tissues including the anterior and posterior midguts, hindgut, and Malpighian tubules. In late embryogenesis, expression is observed in the gonads. In third instar larvae, dpa expression is observed in imaginal tissues. In the eye disc, transcripts are found ahead of the morphogenetic furrow and in a tight band behind it but not in the furrow itself, coinciding with the first and second waves of proliferating cells. In the larval CNS, expression is evident in the optic lobes and in neuroblasts that are producing ganglion mother cells.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
part_of CMG complex
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\dpa in JBrowse
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
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 9 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of dpa
Transgenic constructs containing regulatory region of dpa
Aberrations (Deficiencies and Duplications) ( 43 )
Inferred from experimentation ( 43 )
Inferred from location ( 5 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (9)
14 of 14
Yes
Yes
4 of 14
No
No
3 of 14
No
No
1  
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
2 of 14
No
No
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (9)
14 of 14
Yes
Yes
4 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
Yes
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
14 of 14
Yes
Yes
4 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (12)
11 of 13
Yes
Yes
3 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
Yes
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
Danio rerio (Zebrafish) (10)
14 of 14
Yes
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
2 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (6)
14 of 14
Yes
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
Anopheles gambiae (African malaria mosquito) (10)
12 of 12
Yes
Yes
3 of 12
No
No
Arabidopsis thaliana (thale-cress) (8)
13 of 13
Yes
Yes
4 of 13
No
No
3 of 13
No
No
3 of 13
No
No
3 of 13
No
No
3 of 13
No
No
3 of 13
No
Yes
3 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (6)
12 of 13
Yes
Yes
3 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
2 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (6)
11 of 12
Yes
Yes
3 of 12
No
No
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
Escherichia coli (enterobacterium) (1)
1 of 11
Yes
No
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:dpa. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (7)
7 of 13
6 of 13
6 of 13
6 of 13
6 of 13
6 of 13
1 of 13
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 ( 2 )
    Modifiers Based on Experimental Evidence ( 2 )
    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.
    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
    Interaction Browsers

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    Interaction Browsers

    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
    Subunit Structure (UniProtKB)
    Component of the Mcm2-7 complex. The complex forms a toroidal hexameric ring with the proposed subunit order Mcm2-Mcm6-Mcm4-Mcm7-Mcm3-Mcm5 (Probable).
    (UniProt, Q26454 )
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-56
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    43C6-43C7
    Limits computationally determined from genome sequence between P{lacW}cosk16101 and P{EP}CG1600EP398
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    43C-43C
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (9)
    Genomic Clones (14)
     

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

    cDNA Clones (68)
     

    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
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
       

      polyclonal

      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          The Sld5, Psf1, Psf2, Psf3, CDC45L, Mcm2, Mcm3, Mcm5, Mcm6, Mcm7 and dpa proteins form the Cdc45/Mcm2-7/GINS (CMG) complex, a helicase required for the S phase transition in Kc cells.

          Candidate gene for tibia length quantitative trait locus.

          dpa is misexpressed in the PNS of da mutant embryos.

          dpa is a member of a gene family with extensive sequence identity to the MCM (minichromosome maintenance) genes of S.cerevisiae.

          Physical interactions exist between three MCM proteins, dpa, Mcm2 and Mcm5. The three proteins reside predominantly in stable multi-protein complexes of about 600kD. Coimmunoprecipitation experiments indicate there are at least two distinct 600kD complexes, one of which includes Mcm5 and the other dpa.

          dpa is required for the regulation of DNA replication in neuroblasts and for DNA replication in mitotically proliferating cells.

          Relationship to Other Genes
          Source for database merge of
          Additional comments
          Nomenclature History
          Source for database identify of

          Source for identity of: dpa CG1616

          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (16)
          Reported As
          Symbol Synonym
          Name Synonyms
          Disc proliferation abnormal
          Drosophila proliferation abnormal
          Secondary FlyBase IDs
          • FBgn0004127
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 59 )
          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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
          UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
          Other crossreferences
          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
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          PDB - An information portal to biological macromolecular structures
          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
          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
          KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (132)