FB2025_04 , released October 2, 2025
Gene: Dmel\mod
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General Information
Symbol
Dmel\mod
Species
D. melanogaster
Name
modulo
Annotation Symbol
CG2050
Feature Type
FlyBase ID
FBgn0002780
Gene Model Status
Stock Availability
Gene Summary
modulo (mod) encodes a nucleolar protein that is required for growth of proliferative cells through its association with the proto-oncogene Myc. The mod protein is essential for transcription of spermatid-differentiation genes and for high expression of meiotic arrest genes. The product of mod is phosphorylated, which in turn regulates its localization. [Date last reviewed: 2018-09-20] (FlyBase Gene Snapshot)
Also Known As

ms(3)100EF

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-103
RefSeq locus
NT_033777 REGION:32052060..32054974
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (12 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables DNA binding
inferred from direct assay
inferred from physical interaction with FLYBASE:nej; FB:FBgn0261617
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables mRNA binding
inferred from biological aspect of ancestor with PANTHER:PTN009063129
inferred from electronic annotation with InterPro:IPR035979
enables RNA binding
inferred from electronic annotation with InterPro:IPR000504
Biological Process (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
Cellular Component (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
located_in nucleolus
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from physical interaction with FLYBASE:nej; FB:FBgn0261617
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nuclear speck
inferred from biological aspect of ancestor with PANTHER:PTN000567445
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
Gene Snapshot
modulo (mod) encodes a nucleolar protein that is required for growth of proliferative cells through its association with the proto-oncogene Myc. The mod protein is essential for transcription of spermatid-differentiation genes and for high expression of meiotic arrest genes. The product of mod is phosphorylated, which in turn regulates its localization. [Date last reviewed: 2018-09-20]
Protein Function (UniProtKB)
Its capacity to bind DNA and protein(s), and its differential expression during development suggest a role in the regulation of gene expression during Drosophila development. It could, in interaction with other factors, be required for the translation of instructions provided by pattern forming genes and controls, via chromatin changes, the activity of genes critical for the process of morphogenesis of several embryonic territories.
(UniProt, P13469)
Summary (Interactive Fly)

RRM-containing domain - modifier of PEV promoting chromatin compaction and inactivation - controls cellular growth rate downstream of dMYC - transcriptional regulation by Modulo integrates meiosis and spermatid differentiation in male germ line - the nucleolar protein, Modulo, in complex with CAL1, is essential for the centromeric deposition of the centromere-specific histone H3 variant, CID

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

Please see the JBrowse view of Dmel\mod 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 P13469)

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 5.47

Gene model reviewed during 5.42

Gene model reviewed during 5.53

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0085874
2287
542
FBtr0310268
1954
542
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
UniProt
RefSeq ID
GenBank
FBpp0085233
60.3
542
5.07
FBpp0301951
60.3
542
5.07
Polypeptides with Identical Sequences

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

542 aa isoforms: mod-PA, mod-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Post Translational Modification

The N-terminus is blocked.

(UniProt, P13469)
Crossreferences
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\mod 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).

-0.69

Transcript Expression
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

mod protein is most abundant in embryos but is detected at all developmental stages tested. It is first detected before cellularization in all somatic nuclei, precisely when pericentric heterochromatin becomes visible. After the first cell division, mod protein is found in lineages of specific embryonic primordia. These lineages arise from mitotic domains 1, 2, 4-6, 10, 13, 22, and 25.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
located_in nucleolus
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from physical interaction with FLYBASE:nej; FB:FBgn0261617
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\mod 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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 18 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 17 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of mod
Transgenic constructs containing regulatory region of mod
Aberrations (Deficiencies and Duplications) ( 13 )
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
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
adult thorax & macrochaeta | somatic clone | cell autonomous
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (21)
1 of 14
Yes
No
3  
1 of 14
Yes
No
5  
1 of 14
Yes
No
2  
1 of 14
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No
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No
4  
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No
2  
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No
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3  
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No
1 of 14
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No
20  
1 of 14
Yes
No
1  
1 of 14
Yes
No
1 of 14
Yes
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (18)
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
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No
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No
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No
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No
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No
1 of 14
Yes
No
1 of 14
Yes
No
Mus musculus (laboratory mouse) (16)
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
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No
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No
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No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
Xenopus tropicalis (Western clawed frog) (10)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
Danio rerio (Zebrafish) (20)
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
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Yes
No
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No
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No
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No
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No
1 of 14
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No
Caenorhabditis elegans (Nematode, roundworm) (8)
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
Anopheles gambiae (African malaria mosquito) (18)
Arabidopsis thaliana (thale-cress) (39)
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
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No
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Yes
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Yes
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Yes
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No
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No
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No
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No
1 of 13
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (2)
1 of 13
Yes
No
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (4)
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
Yes
1 of 12
Yes
Yes
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:mod. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (29)
3 of 13
2 of 13
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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 ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    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?
    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

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3R
    Recombination map
    3-103
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    100F2-100F2
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    100F5-100F5
    100E1-100F5
    (determined by in situ hybridisation)
    100F-100F
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (13)
    Genomic Clones (11)
     

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

    cDNA Clones (326)
     

    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)
      Other clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          The size of the mod protein differs between testes and somatic tissues. The testes-specific mod variant has an apparent molecular mass that is consistent with the predicted 60.3 kDa of the full size protein. The mod variant expressed in somatic tissues has an apparent molecular mass of approximately 50 kDa and is missing the acidic N-terminal domain found in the full size protein.

          Both the full-size testes-specific variant and the 50 kDa somatic variant of the mod protein specifically bind to the conserved testis-specific TSE promoter motif.

          mod is required selectively in proliferative cells to sustain their growth and maintain their specific size.

          mod is a genetic target of tsh. A biological role of tsh is to repress mod expression in the ectoderm and this negative control is performed independently of Scr. tsh protein binds to specific DNA sequences within a 5' mod regulatory region that reproduces tsh-dependent expression in the ectoderm.

          mod gene acts downstream of pattern forming genes of both the embryonic axes and that its function is required for an accurate morphogenesis of several tissue types.

          Postmeiotic differentiation defect.

          mod is a haplo suppressor of position effect variegation (PEV). At the genetic level mod acts downstream of Ubx, Scr, tsh, dl, twi and sna. Mutant analysis demonstrates mod is required to allow accurate morphogenesis to occur. Together these results suggest that mod could act as a 'memory' of selector gene information regulating the expression of the gene directly involved in morphogenesis via chromatin structure.

          Mutant phenotype, expression pattern during embryogenesis and DNA binding activity of gene product suggest that mod regulates chromatin structure and activity in specific cell lineages.

          P element transformation experiments demonstrated that mod+ can replace the function of Map205.

          Relationship to Other Genes
          Source for database merge of

          Source for merge of: mod ms(3)100EF

          Additional comments

          "ird6" may correspond to the "krz" or "mod" gene or both; the ird61 mutation fails to complement the lethality of "krz" alleles (krzS047819) and deficiences which affect "mod" (Df(3R)A4-4L2 and Df(3R)A4-4L3).

          "ird6" stated to correspond to "mod".

          may be allelic to e(gs)

          Nomenclature History
          Source for database identify of

          Source for identity of: mod CG2050

          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (8)
          Reported As
          Secondary FlyBase IDs
          • FBgn0011255
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 55 )
          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)
          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.
          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
          References (162)