FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\His2A
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General Information
Symbol
Dmel\His2A
Species
D. melanogaster
Name
Histone H2A
Annotation Symbol
Feature Type
FlyBase ID
FBgn0001196
Gene Model Status
Stock Availability
Gene Summary
Core component of nucleosome. Nucleosomes wrap and compact DNA into chromatin, limiting DNA accessibility to the cellular machineries which require DNA as a template. Histones thereby play a central role in transcription regulation, DNA repair, DNA replication and chromosomal stability. DNA accessibility is regulated via a complex set of post-translational modifications of histones, also called histone code, and nucleosome remodeling. Involved in recruitment of Parp1 to chromatin and regulates its activity; the N-terminal tail inhibits histone H4-dependent activation of Parp1 (PubMed:17827147, PubMed:24508391). (UniProt, P84051)
Contribute a Gene Snapshot for this gene.
Also Known As

H2A, histone, core histone, HisC, dH2A

Function
Gene Ontology (GO) Annotations (12 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables DNA binding
inferred from electronic annotation with InterPro:IPR002119, InterPro:IPR007125
inferred from sequence or structural similarity with UniProtKB:Q7L7L0
non-traceable author statement
inferred from electronic annotation with InterPro:IPR002119
inferred from biological aspect of ancestor with PANTHER:PTN000604564
Biological Process (4 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from mutant phenotype
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with MGI:MGI:2448285
inferred from biological aspect of ancestor with PANTHER:PTN000604564
non-traceable author statement
Cellular Component (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
located_in chromosome
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
part_of nucleosome
non-traceable author statement
inferred from electronic annotation with InterPro:IPR002119
traceable author statement
inferred from sequence or structural similarity with UniProtKB:Q7L7L0
inferred from biological aspect of ancestor with PANTHER:PTN000604564
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000604564
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the histone H2A family. (P84051)
Summaries
Protein Function (UniProtKB)
Core component of nucleosome. Nucleosomes wrap and compact DNA into chromatin, limiting DNA accessibility to the cellular machineries which require DNA as a template. Histones thereby play a central role in transcription regulation, DNA repair, DNA replication and chromosomal stability. DNA accessibility is regulated via a complex set of post-translational modifications of histones, also called histone code, and nucleosome remodeling. Involved in recruitment of Parp1 to chromatin and regulates its activity; the N-terminal tail inhibits histone H4-dependent activation of Parp1 (PubMed:17827147, PubMed:24508391).
(UniProt, P84051)
Summary (Interactive Fly)

Chromatin component - a core histone that can be modified by ubiquitination - a target of polycomb gene silencing - involved in trans-histone regulation through histone H3 - H2A monoubiquitination promotes histone H3 methylation in Polycomb repression

Gene Model and Products
Number of Transcripts
0
Number of Unique Polypeptides
0
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 P84051)

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
Transcript Data
Annotated Transcripts
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Polypeptides with Identical Sequences

 

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

The nucleosome is a histone octamer containing two molecules each of H2A, H2B, H3 and H4 assembled in one H3-H4 heterotetramer and two H2A-H2B heterodimers. The octamer wraps approximately 147 bp of DNA. Interacts with Parp1 (via C-terminus); the interaction is direct and regulates Parp1 enzymatic activity (PubMed:17827147).

(UniProt, P84051)
Post Translational Modification

The chromatin-associated form, but not the free cytoplasmic form, is phosphorylated on Thr-120 by NHK-1 during mitosis, and dephosphorylated during S-phase. Also phosphorylated on Thr-120 by NHK-1 during prophase I of meiosis; which is required for acetylation of H3 'Lys-14' and H4 'Lys-5', disassembly of the synaptonemal complex, and karyosome formation.

Monoubiquitination of Lys-119 by sce/dRING gives a specific tag for epigenetic transcriptional repression (PubMed:15386022). Deubiquitinated by the polycomb repressive deubiquitinase (PR-DUB) complex (PubMed:20436459, PubMed:30258054, PubMed:30639226).

Phosphorylation on Ser-2 is enhanced during mitosis. Phosphorylation on Ser-2 directly represses transcription (By similarity).

(UniProt, P84051)
Crossreferences
PDB - An information portal to biological macromolecular structures
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\His2A 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).

NA

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

His2A transcript is restricted to replicating cells.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

His4 and His2A that was mono or poly acetylated is detected associated with nuclei of developing spermatids until late stages. During late spermatid development and in mature spermatazoa these proteins can no longer be detected (and are presumably replaced by sperm specific chromatin packaging proteins). However, shortly after fertilization His4 and His2A immunoreactivity reappears associated with the male pronuclear DNA.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in chromosome
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\His2A 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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 1 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 38 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of His2A
Transgenic constructs containing regulatory region of His2A
Aberrations (Deficiencies and Duplications) ( 5 )
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
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (0)
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (0)
Mus musculus (laboratory mouse) (0)
Xenopus tropicalis (Western clawed frog) (0)
Danio rerio (Zebrafish) (0)
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (0)
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:His2A. Refer to their site for version information.
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 ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    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)
    The nucleosome is a histone octamer containing two molecules each of H2A, H2B, H3 and H4 assembled in one H3-H4 heterotetramer and two H2A-H2B heterodimers. The octamer wraps approximately 147 bp of DNA. Interacts with Parp1 (via C-terminus); the interaction is direct and regulates Parp1 enzymatic activity (PubMed:17827147).
    (UniProt, P84051 )
    Linkouts
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    Recombination map
    2-55
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    39D3-39E1
    Left limit from in situ hybridisation (FBrf0029738) Right limit from inclusion within Df(2L)L138D-XR (citation unavailable)
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    39D-39E
    (determined by in situ hybridisation)
    39D3-39E2
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location

    2-55

    Left of (cM)
    Right of (cM)
    Notes

    2-54.6

    Stocks and Reagents
    Stocks (30)
    Genomic Clones (0)
     
      cDNA Clones (0)
       

      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
                Antibody Information
                Laboratory Generated Antibodies
                Commercially Available Antibodies
                 
                Cell Line Information
                Publicly Available Cell Lines
                 
                  Other Stable Cell Lines
                   
                    Other Comments

                    The ATPase activity of Iswi is completely inhibited by each of the four histone tails (His2A, His2B, His3 and His4), results indicate a novel role for the flexible histone tails in chromatin remodeling by Iswi.

                    The codon bias of the histone genes from D.melanogaster and D.hydei illustrates that the generalisation that abundantly expressed genes have a high codon bias and low rates of silent substitution does not hold for the histone genes.

                    The position of the homologous histone gene repeats within the nuclei of early embryo cells has been investigated. The two homologous histone gene clusters are distinct and separate through all stages of the cell cycle up to nuclear cycle 13. During interphase of cycle 14, the two clusters colocalise with high frequency, and move from near the midline of the nucleus towards the apical side.

                    DNA replication of the 5kb histone gene repeating unit in tissue culture cells (Drosophila Kc cells) initiates at multiple sites located within the repeating unit. Several replication pause sites are located at 5' upstream regions of some histone genes.

                    DNaseI footprinting analysis reveals core histones His2A, His2B, His3 and His4 (but not His1) bind to the kni, Kr and Ubx minimal enhancer elements in a periodic manner.

                    The genomic organisation of the histone genes in D.hydei closely resembles that of D.melanogaster.

                    One H2A-like sequence variant has been found (Donahue, Palmer, Condie, Sabatini, and Blumenfeld, 1986); see His2Av.

                    Encodes Histone-2A. See HIS-C record.

                    Relationship to Other Genes
                    Nomenclature History
                    Source for database identify of
                    Nomenclature comments
                    Etymology
                    Synonyms and Secondary IDs (23)
                    Reported As
                    Symbol Synonym
                    H2A
                    (Karling and Weavers, 2025, Pérez-Roldán et al., 2025, Gilbert et al., 2024, Hunt and Mannervik, 2024, Kurshakova et al., 2024, Parreno et al., 2024, Zhang et al., 2024, Zhao et al., 2024, Corcoran and Jacob, 2023, Dubruille et al., 2023, Erokhin et al., 2023, Hodkinson et al., 2023, Tirgar et al., 2023, Zhang et al., 2023, Bonnet et al., 2022, Ranjan and Chen, 2022, Urban et al., 2022, Chaouch and Lasko, 2021, Cho et al., 2021, Endo et al., 2021, Khan et al., 2021, Llorens-Giralt et al., 2021, Matsuo, 2021, Okimune et al., 2021, Stephenson et al., 2021, Tilly et al., 2021, Zhang et al., 2021, Barbour et al., 2020, Kolesnikova et al., 2020, Kuroda et al., 2020, Morgan and Shilatifard, 2020, Okimune et al., 2020, Pirani et al., 2020, Tarczewska et al., 2020, Wooten et al., 2020, Chittori et al., 2019, Wooten et al., 2019, Yang et al., 2019, Armstrong et al., 2018, Daou et al., 2018, Tsui et al., 2018, Zheng et al., 2018, Janssens et al., 2017, Khuong et al., 2017, Kitevski-LeBlanc et al., 2017, Kolkhof et al., 2017, Ramachandran et al., 2017, Rieder et al., 2017, Rowley et al., 2017, Strom et al., 2017, Yang and Ioshikhes, 2017, Bayona-Feliu et al., 2016, Doiguchi et al., 2016, Elnfati et al., 2016, Kahn et al., 2016, Kavi et al., 2016, Messina et al., 2016, Nakashima et al., 2016, Ozawa et al., 2016, Penke et al., 2016, Shih et al., 2016, Zhou et al., 2016, Edlich-Muth et al., 2015, Fei et al., 2015, Horard and Loppin, 2015, Pengelly et al., 2015, Emelyanov et al., 2014, Fereres et al., 2014, Klinker et al., 2014, Kusch et al., 2014, Landais et al., 2014, Messina et al., 2014, Thomas et al., 2014, Bharadwaj et al., 2013, Doyen et al., 2013, Eichinger et al., 2013, Guglielmi et al., 2013, Schaaf et al., 2013, Stein et al., 2013, Wang et al., 2013, Xing et al., 2013, Zhou et al., 2013, Dunlap et al., 2012, Gutiérrez et al., 2012, Ito et al., 2012, Li et al., 2012, Li et al., 2012, Lo et al., 2012, Villar-Garea et al., 2012, Xie et al., 2012, Zhou et al., 2012, Chen et al., 2011, Egelhofer et al., 2011, Gibert and Karch, 2011, Lorbeck et al., 2011, Olins et al., 2011, Regnard et al., 2011, Siudeja et al., 2011, Anderson et al., 2010, Beck et al., 2010, Bryson et al., 2010, Deal et al., 2010, Guertin and Lis, 2010, Lancaster et al., 2010, Makde et al., 2010, Sawatsubashi et al., 2010, Scheuermann et al., 2010, Weber et al., 2010, Kolesnikova et al., 2009, Lu et al., 2009, Morciano et al., 2009, Sakai et al., 2009, Sullivan et al., 2009, Bao et al., 2008, Barreau et al., 2008, Cakouros et al., 2008, Clapier et al., 2008, Frydrychova et al., 2008, Hanai et al., 2008, Krajewski, 2008, Lagarou et al., 2008, Bell et al., 2007, Camporeale et al., 2007, Corona et al., 2007, Dalal et al., 2007, Dalal et al., 2007, Isogai et al., 2007, Pinnola et al., 2007, Rathke et al., 2007, Tie et al., 2007, Wagner et al., 2007, Wagner et al., 2007, Brasaemle and Hansen, 2006, Camporeale et al., 2006, Furuyama et al., 2006, Mendjan et al., 2006, Raisner and Madhani, 2006, Santoso and Kadonaga, 2006, Dudnik et al., 2005, Ivanovska et al., 2005, Jin, 2005, Lusser et al., 2005, Aihara et al., 2004, Kusch et al., 2004, Kusch et al., 2004, Lippman and Martienssen, 2004, Morales et al., 2004, Wang et al., 2004, Furukawa et al., 2003, Kakita et al., 2003, Schwartz et al., 2003, Staeva-Vieira et al., 2003, Blower and Karpen, 2002, Cao et al., 2002, Clapier et al., 2002, Levenstein and Kadonaga, 2002, Smith, 2002, Yu and Wolfner, 2002, Berloco et al., 2001, Hamiche et al., 2001, Katsani et al., 2001, Loppin et al., 2001, Mello and Almouzni, 2001, Mizuguchi et al., 2001, Nakagawa et al., 2001, Kal et al., 2000, Leach et al., 2000, Mizzen and Allis, 2000, Pham and Sauer, 2000, Smith et al., 2000, Verreault, 2000, Baldo et al., 1999, Borgnetto et al., 1999, Carrier et al., 1999, Hamiche et al., 1999, Reim et al., 1999, Stuckenholz et al., 1999, Schienman et al., 1998, Sobel et al., 1998, Strausbaugh et al., 1998, Walker and Bownes, 1998, Georgel et al., 1997, Grunstein, 1997, Ito et al., 1996, Ito et al., 1996, Sommer and Strausbaugh, 1996, Strausbaugh and Williams, 1996, Wolffe and Pruss, 1996, Blank and Becker, 1995, Sobel et al., 1994, O'Brien and Lis, 1993, Shinomiya and Ina, 1993, Becker and Wu, 1992, Kas and Laemmli, 1992, Kerrigan and Kadonaga, 1992, Pauli et al., 1992, Harisanova and Ralchev, 1991, Harisanova et al., 1991, O'Brien and Lis, 1991, Udvardy and Schedl, 1991, Fitch et al., 1990, Kremer and Hennig, 1990, Domier et al., 1986, Levinger and Varshavsky, 1982)
                    Secondary FlyBase IDs
                      Datasets (0)
                      Study focus (0)
                      Experimental Role
                      Project
                      Project Type
                      Title
                      Study result (0)
                      Result
                      Result Type
                      Title
                      External Crossreferences and Linkouts ( 33 )
                      Sequence Crossreferences
                      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.
                      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.
                      PDB - An information portal to biological macromolecular structures
                      Linkouts
                      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                      References (415)