FB2025_05 , released December 11, 2025
Gene: Dmel\Clamp
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General Information
Symbol
Dmel\Clamp
Species
D. melanogaster
Name
Chromatin-linked adaptor for MSL proteins
Annotation Symbol
CG1832
Feature Type
FlyBase ID
FBgn0032979
Gene Model Status
Stock Availability
Gene Summary
Chromatin-linked adaptor for MSL proteins (Clamp) encodes a zinc finger protein that is required to recruit the dosage compensation complex. It is essential in both males and females. It also functions outside of the dosage compensation pathway. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-54
RefSeq locus
NT_033779 REGION:22165720..22169143
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (15 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
is_active_in chromatin
inferred from direct assay
located_in chromatin
inferred from direct assay
located_in chromosome
inferred from direct assay
inferred from mutant phenotype
is_active_in histone locus body
inferred from direct assay
located_in nucleus
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
Protein Family (UniProt)
-
Summaries
Gene Snapshot
Chromatin-linked adaptor for MSL proteins (Clamp) encodes a zinc finger protein that is required to recruit the dosage compensation complex. It is essential in both males and females. It also functions outside of the dosage compensation pathway. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
C2H2 ZINC FINGER TRANSCRIPTION FACTORS -
Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys2His2 zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).
Protein Function (UniProtKB)
Transcription factor involved in X-chromosome dosage compensation in males, the process by which transcription of the single X chromosome in the male is elevated (PubMed:23873939, PubMed:27414415, PubMed:27995349, PubMed:31320325, PubMed:34187599, PubMed:37602401). Binds to the DNA sequence (GA)n (PubMed:27414415, PubMed:28838946, PubMed:29281702). Clamp-binding promotes nucleosome depletion and chromatin accessibility, thereby allowing access to other transcription factors (PubMed:28838946, PubMed:29077765, PubMed:34187599). Specifically binds to cis-acting elements on the X-chromosome named chromatin entry sites and promotes recruitment of the male-specific lethal (MSL) histone acetyltransferase complex, which associates with actively transcribed genes on the male X-chromosome to upregulate their expression (PubMed:23873939, PubMed:27414415, PubMed:31320325). Mechanistically, acts by promoting chromatin accessibility at chromatin entry sites, facilitating DNA-binding of msl-2, followed by MSL complex recruitment (PubMed:31320325, PubMed:37602401). In addition to dosage compensation, also involved in zygotic genome activation (ZGA), a critical event in early embryonic development during which the developmental control passes from maternally provided mRNAs to the expression of the zygotic genome after fertilization (PubMed:34342574, PubMed:34849887). Maternally-provided protein cooperates with Zelda (zld) to activate zygotic transcription by increasing chromatin accessibility at promoters of specific genes and facilitate zld occupancy at a subset of key embryonic promoters (PubMed:34342574). Also acts as an activator of gypsy chromatin insulator function by promoting binding of Cp190 to chromatin (PubMed:30718365).
(UniProt, Q9V9N4)
Summary (Interactive Fly)

zinc finger transcription factor - dosage compensation - essential in both males and females - also functions outside of the dosage compensation pathway

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

Please see the JBrowse view of Dmel\Clamp 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 Q9V9N4)

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

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

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

Gene model reviewed during 5.45

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0085964
2636
561
FBtr0100407
2897
566
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
FBpp0085317
61.2
561
9.06
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Homodimer (PubMed:35409222). Interacts with msl-2; promoting recruitment of the male-specific lethal (MSL) histone acetyltransferase complex to chromatin (PubMed:31320325, PubMed:35648444, PubMed:37602401). Interacts with Nelf-A (PubMed:29281702). Interacts with NELF-B (PubMed:29281702).

(UniProt, Q9V9N4)
Crossreferences
InterPro - A database of protein families, domains and functional sites
PDB - An information portal to biological macromolecular structures
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\Clamp using the Feature Mapper tool.

External Data
Crossreferences
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).

0.78

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

Comment: maternally deposited

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
is_active_in chromatin
inferred from direct assay
located_in chromatin
inferred from direct assay
located_in chromosome
inferred from direct assay
inferred from mutant phenotype
is_active_in histone locus body
inferred from direct assay
located_in nucleus
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Clamp 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) 1-3
  • Stages(s) 4-6
  • Stages(s) 13-16
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 6 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 21 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Clamp
Transgenic constructs containing regulatory region of Clamp
Aberrations (Deficiencies and Duplications) ( 0 )
Inferred from experimentation ( 0 )
Inferred from location ( 2 )
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) (290)
2 of 14
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Hsap\LOC344065
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No
Hsap\LOC728743
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Yes
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Hsap\ZNF660-ZNF197
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Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (84)
2 of 14
Yes
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
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No
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Yes
Mus musculus (laboratory mouse) (135)
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No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
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Yes
1 of 14
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No
1 of 14
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Yes
1 of 14
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No
1 of 14
No
Yes
1 of 14
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Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (68)
2 of 13
Yes
Yes
2 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (62)
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
Yes
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
Yes
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
Yes
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
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
Yes
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
Yes
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
Yes
1 of 14
Yes
No
1 of 14
Yes
Yes
1 of 14
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (9)
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
1 of 14
Yes
No
Anopheles gambiae (African malaria mosquito) (46)
10 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (13)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
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
Saccharomyces cerevisiae (Brewer's yeast) (3)
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Clamp. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (62)
2 of 13
2 of 13
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2 of 13
2 of 13
2 of 13
1 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
    Interaction Browsers

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    RNA-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    Interaction Browsers
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Subunit Structure (UniProtKB)
    Homodimer (PubMed:35409222). Interacts with msl-2; promoting recruitment of the male-specific lethal (MSL) histone acetyltransferase complex to chromatin (PubMed:31320325, PubMed:35648444, PubMed:37602401). Interacts with Nelf-A (PubMed:29281702). Interacts with NELF-B (PubMed:29281702).
    (UniProt, Q9V9N4 )
    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
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map
    2-54
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    40F2-40F2
    Limits computationally determined from genome sequence between P{lacW}Df31k05815 and P{PZ}cul-202074&P{lacW}l(2)k08110k08110
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (15)
    Genomic Clones (23)
    cDNA Clones (106)
     

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

          Source for identity of: Clamp CG1832

          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (6)
          Reported As
          Symbol Synonym
          Name Synonyms
          Chromatin-linked adaptor for MSL proteins
          chromatin-linked adaptor for MSL proteins
          Secondary FlyBase IDs
          • FBan0001832
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 39 )
          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
          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)
          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
          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
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (109)