FlyBase Gene Report: Dmel\His2Av

General Information

Symbol

Dmel\His2Av

Species

D. melanogaster

Name

Histone H2A variant

Annotation Symbol

CG5499

Feature Type

protein_coding_gene

FlyBase ID

FBgn0001197

Gene Model Status

Current

Stock Availability

42 publicly available

Gene Summary

Histone H2A variant (His2Av) encodes a protein that is present in about 25% of nucleosomes. It influences chromatin structure and gene regulation and is also phosphorylated in response to DNA damage in the less conserved C-terminal tail. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)

All Summaries

Also Known As

H2Av, H2AvD, His2AvD, H2A.Z, γ-H2Av

Key Links

Genomic Location

Cytogenetic map

97D3-97D3

Sequence location

3R:26,866,929..26,869,220 [+]

Recombination map

3-92

RefSeq locus

NT_033777 REGION:26866929..26869220

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (18 terms)

Molecular Function (4 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables protein-containing complex binding

inferred from direct assay

(Kusch et al., 2004)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

enables DNA binding

inferred from electronic annotation with InterPro:IPR002119, InterPro:IPR007125

(InterPro Project Members, 2004-)

enables protein heterodimerization activity

inferred from electronic annotation with InterPro:IPR009072

(InterPro Project Members, 2004-)

enables structural constituent of chromatin

inferred from electronic annotation with InterPro:IPR002119

(InterPro Project Members, 2004-)

inferred from biological aspect of ancestor with PANTHER:PTN000604564

(GO Reference Genome Project, 2011-)

Biological Process (7 terms)

Terms Based on Experimental Evidence (6 terms)

CV Term

Evidence

References

involved_in histone locus body assembly

inferred from mutant phenotype

(Wagner et al., 2007)

involved_in male germ-line stem cell population maintenance

inferred from mutant phenotype

(Morillo Prado et al., 2013)

involved_in negative regulation of peptidoglycan recognition protein signaling pathway

inferred from genetic interaction with FLYBASE:Su(var)2-10; FB:FBgn0003612,FLYBASE:Rel; FB:FBgn0014018

(Tang et al., 2021)

involved_in protein localization to chromatin

inferred from direct assay

(Kotova et al., 2011)

involved_in protein localization to nuclear body

inferred from mutant phenotype

(Wagner et al., 2007)

involved_in somatic stem cell population maintenance

inferred from mutant phenotype

(Morillo Prado et al., 2013)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

involved_in heterochromatin formation

inferred from biological aspect of ancestor with PANTHER:PTN000604564

(GO Reference Genome Project, 2011-)

Cellular Component (7 terms)

Terms Based on Experimental Evidence (7 terms)

CV Term

Evidence

References

located_in chromosome

inferred from direct assay

(Leach et al., 2000)

located_in euchromatin

inferred from mutant phenotype

(Scacchetti et al., 2020)

part_of nucleosome

inferred from direct assay

(Mavrich et al., 2008)

located_in nucleus

inferred from direct assay

(Leach et al., 2000)

located_in polytene chromosome chromocenter

inferred from direct assay

(Andreyeva et al., 2008)

located_in polytene chromosome

inferred from direct assay

(Andreyenkova et al., 2009, Leach et al., 2000)

located_in site of double-strand break

inferred from direct assay

(Lake et al., 2015)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

part_of nucleosome

inferred from biological aspect of ancestor with PANTHER:PTN000604564

(GO Reference Genome Project, 2011-)

inferred from electronic annotation with InterPro:IPR002119

(InterPro Project Members, 2004-)

is_active_in nucleus

inferred from biological aspect of ancestor with PANTHER:PTN000604564

(GO Reference Genome Project, 2011-)

Gene Group (FlyBase)

Pathway (FlyBase)

NEGATIVE REGULATORS OF IMD SIGNALING PATHWAY

Protein Family (UniProt)

Belongs to the histone H2A family. (P08985)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

Pathway (FlyBase)

NEGATIVE REGULATORS OF IMD SIGNALING PATHWAY -: Negative regulators of the immune deficiency (Imd) pathway result in the decreased activity of the NFκB-like transcription factor Rel in the nucleus. Negative regulators are important in preventing damage to the host from over-activation of the pathway; preventing inappropriate triggering or terminating the response. (Adapted from FBrf0224587 and FBrf0238555.)

Protein Function (UniProtKB)

Variant histone H2A which replaces conventional H2A in a subset of nucleosomes (PubMed:1498368). Nucleosomes wrap and compact DNA into chromatin, limiting DNA accessibility to the cellular machineries which require DNA as a template (Probable). Histones thereby play a central role in transcription regulation, DNA repair, DNA replication and chromosomal stability (Probable). DNA accessibility is regulated via a complex set of post-translational modifications of histones, also called histone code, and nucleosome remodeling (Probable). Acts as a Polycomb group (PcG) protein required to maintain the transcriptionally repressive state of homeotic genes of the animal throughout development (PubMed:15630020). Required for histone H3 'Lys-9' methylation and histone H4 'Lys-12' acetylation, two modifications that are essential for heterochromatin formation (PubMed:15630020). Involved in the chromatin recruitment of Parp1 and the regulation of its ADP-ribosylation activity to alter the local chromatin state in response to various stimuli, including heat shock and genotoxic stress (PubMed:21444826). Also involved in DNA double strand break (DSB) repair (PubMed:12202754). Essential for early development (PubMed:10385122).

(UniProt, P08985)

Summary (Interactive Fly)

replacement histone participates in an ordered cascade of events leading to the establishment of heterochromatin - controls poly(ADP-ribose) polymerase 1 (PARP1) activation in chromatin

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\His2Av 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 P08985)

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

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

His2Av-RA

FBtr0085062

NM_079795

912

141

His2Av-RB

FBtr0334311

NM_001276068

1169

141

Additional Transcript Data and Comments

Reported size (kB)

0.9 (northern blot)

(van Daal and Elgin, 1992)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

His2Av-PA

FBpp0084434

15.0

141

10.94

FBpp0306426

15.0

141

10.94

P08985

NP_001262997

AGB96377

Polypeptides with Identical Sequences

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

141 aa isoforms: His2Av-PA, His2Av-PB

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. H2A or its variant His2Av forms a heterodimer with H2B (Probable). Interacts with Nasp; this interaction directly or indirectly destabilizes His2Av (PubMed:36930688). Interacts with the kinase JIL-1 (PubMed:24508391).

(UniProt, P08985)

Post Translational Modification

Phosphorylated on Ser-138 by the RPS6KA5 ortholog kinase JIL-1 (PubMed:12202754, PubMed:21444826, PubMed:24508391). Phosphorylation of Ser-138 leads to activation of Parp1 triggering Parp1-dependent chromatin processes, including chromatin remodeling induced by poly-ADP-ribosylation of Parp1 and other chromatin proteins; phosphorylation at Ser-138 may alter nucleosomal organization to expose the H4 C-terminal tail, which interacts with and activates Parp1 (PubMed:21444826, PubMed:24508391). Phosphorylation is required for the genotoxic stress response (PubMed:21444826). Phosphorylation of Ser-138 in response to DNA double strand breaks (DSBs) generated by exogenous genotoxic agents is dependent on the DNA damage checkpoint kinases ATR and ATM, spreads on either side of a detected DSB site and may mark the surrounding chromatin for recruitment of proteins required for DNA damage signaling and repair (PubMed:12202754).

Acetylated on Lys-5 by Tip60. Acetylation is enhanced by Ser-138 phosphorylation and promotes the exchange of the phosphorylated form with the unmodified form of H2AV.

Monoubiquitination of Lys-121 by sce/dRING gives a specific tag for epigenetic transcriptional repression.

(UniProt, P08985)

Domain

The

(UniProt, P08985)

Crossreferences

InterPro - A database of protein families, domains and functional sites

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

P08985

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\His2Av 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).

-0.36

Transcript Expression

No Assay Recorded

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 15

Comment: reference states 0-12 hr AEL

(van Daal and Elgin, 1992)

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 9 -- 10

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 13 -- 16

embryonic brain

(Fisher et al., 2012)

embryonic central brain neuron

(Fisher et al., 2012)

embryonic dorsal epidermis

(Fisher et al., 2012)

embryonic/larval midgut

(Fisher et al., 2012)

larval nervous system

(Fisher et al., 2012)

sensory nervous system primordium

(Fisher et al., 2012)

larval ventral nerve cord

(Fisher et al., 2012)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage | female

(van Daal and Elgin, 1992)

Additional Descriptive Data

His2Av transcripts are abundant in adult females and in embryos throughout the first 12 hours.

(van Daal and Elgin, 1992)

His2Av⁸¹⁰ RNA accumulates to approximately the same level as the wild type RNA.

(van Daal and Elgin, 1992)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage | female

germarium

(Joyce et al., 2011)

adult stage | female & oogenesis

germarium region 2a

(Hur et al., 2016)

oogenesis stage S1 -- S3

nurse cell

(Joyce et al., 2011)

oocyte

(Joyce et al., 2011)

oogenesis stage S1 -- S5

follicle cell

(Joyce et al., 2011)

mass spectroscopy

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage

adult head

• membrane

(Aradska et al., 2015)

day 5 of adulthood | male -- day 7 of adulthood | male

spermatozoon

(Dorus et al., 2006)

day 7 of adulthood -- day 49 of adulthood

adult heart

(Cammarato et al., 2011)

Additional Descriptive Data

His2Av is specific to region 2a and appears in no other part of the germarium.

(Hur et al., 2016)

His2Av is detected in the germarium and in the nucleus of the oocyte and nurse cells as well as mitotically-dividing follicle cells until oogenesis stage S3. After this, staining is dramatically reduced in the nurse cells and oocyte but persists in the follicle cells.

(Joyce et al., 2011)

The His2Av protein is present at nearly constant levels in all developmental stages tested.

(van Daal and Elgin, 1992)

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in chromosome

inferred from direct assay

(Leach et al., 2000)

located_in euchromatin

inferred from mutant phenotype

(Scacchetti et al., 2020)

part_of nucleosome

inferred from direct assay

(Mavrich et al., 2008)

located_in nucleus

inferred from direct assay

(Leach et al., 2000)

located_in polytene chromosome chromocenter

inferred from direct assay

(Andreyeva et al., 2008)

located_in polytene chromosome

inferred from direct assay

(Andreyenkova et al., 2009, Leach et al., 2000)

located_in site of double-strand break

inferred from direct assay

(Lake et al., 2015)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\His2Av in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\His2Av

View exonic expression by tissue for Dmel\His2Av

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

FBgn0001197

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

43229/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0001197

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0001197

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0001197

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG5499

Flygut - An atlas of the Drosophila adult midgut

FBgn0001197

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\His2Av

Stages(s) 4-6
Stages(s) 7-8
Stages(s) 11-12
Stages(s) 13-16

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 19 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 54 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of His2Av

Transgenic constructs containing regulatory region of His2Av

Aberrations (Deficiencies and Duplications) ( 20 )

Inferred from experimentation ( 20 )

Gene disrupted in

Df(3R)BSC496

(Christensen et al., 2008.4.15)

Df(3R)BSC524

(Christensen et al., 2008.6.11)

Df(3R)ro80b

(Cook and Carpenter, 2002, Kidd et al., 1999)

Df(3R)ro-XB3

(Kidd et al., 1999, Yasuda et al., 1995, Strutt and White, 1994, Knibb et al., 1993)

Df(3R)Tl-I

(Cook and Carpenter, 2002)

Df(3R)Tl-P

(Cook and Carpenter, 2002, Spradling et al., 1999, BDGP Project Members, 1994-1999)

Gene partially disrupted in

Df(3R)nm136

(Martinek et al., 2008)

Gene duplicated in

Dp(3;2)CH322-97107

(Shahab et al., 2015)

Inferred from location ( 6 )

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

cell lethal | somatic clone

His2Av^KO

lethal

lethal, with Scer\GAL4^29BD

lethal, with Scer\GAL4^Act5C.PI

His2Av^HM05177

lethal, with Scer\GAL4^c355

lethal, with Scer\GAL4^en.PU

His2Av^HM05177

lethal | recessive

lethal - all die before end of larval stage

lethal - all die before end of larval stage | recessive

His2Av⁸¹⁰

partially lethal - majority die

His2Av⁰⁵¹⁴⁶

partially lethal - majority die, with Scer\GAL4^hs.2sev

His2Av^GS3127

partially lethal - majority die, with Scer\GAL4^pnr-MD237

His2Av^GD4747

partially lethal - majority die | pupal stage

His2Av⁸¹⁰

partially lethal - majority die | recessive

His2Av⁰⁵¹⁴⁶

partially lethal - majority die | third instar larval stage

His2Av⁸¹⁰

viable

His2Av^72b

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

His2Av^GD4747

viable, with Scer\GAL4^tin.CΔ4

His2Av^GD4747

Sterility

Allele

female semi-fertile, with Scer\GAL4^{VP16.mat.αTub67C}

His2Av^HMS00162

female semi-sterile, with Scer\GAL4^nanos.PG

His2Av^KK108652

female semi-sterile, with Scer\GAL4^{VP16.mat.αTub67C}

His2Av^HM05177

female semi-sterile, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

female sterile, with Scer\GAL4^C587

His2Av^HM05177

female sterile, with Scer\GAL4^nanos.PG

His2Av^GD4747

female sterile, with Scer\GAL4^tj.PU

His2Av^HM05177

Other Phenotypes

Allele

abnormal flight, with Scer\GAL4^Mef2.PR

His2Av^GD4747

abnormal pain response | adult stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

His2Av^GD4747

abnormal size | adult stage, with Scer\GAL4^ey.PH

His2Av^HM05177

decreased cell number | adult stage, with Scer\GAL4^Tk.PS

His2Av^HMS02773

enhancer of variegation, with P{hsp26-pt-T}39C-3

His2Av⁸¹⁰

melanotic mass phenotype | larval stage, with Scer\GAL4^Act5C.PI

His2Av^HM05177

radiation sensitive

His2Av⁸¹⁰

short lived | heat sensitive, with Scer\GAL4^tin.CΔ4

His2Av^GD4747

some die during larval stage

His2Av⁸¹⁰

some die during larval stage | recessive

His2Av⁸¹⁰

some die during pupal stage, with Scer\GAL4^pnr-MD237

His2Av^GD4747

suppressor of variegation, with P{hsp26-pt-T}118E-10

His2Av⁸¹⁰

suppressor of variegation, with P{hsp26-pt-T}gw^39C-33

His2Av⁸¹⁰

suppressor of variegation | dominant

His2Av⁸¹⁰

suppressor of variegation | dominant, with In(1)w^m4

His2Av⁸¹⁰

visible, with Scer\GAL4^dpp.blk1

visible, with Scer\GAL4^hs.2sev

visible, with Scer\GAL4^pnr-MD237

His2Av^GD4747

visible | adult stage, with Scer\GAL4^en.PU

His2Av^HM05177

visible | adult stage, with Scer\GAL4^ey.PH

His2Av^HM05177

visible | adult stage, with Scer\GAL4^GMR.PF

His2Av^HM05177

visible | adult stage, with Scer\GAL4^nub.PK

His2Av^HM05177

visible | somatic clone

His2Av⁸¹⁰

wild-type

Phenotype manifest in

Allele

adult anterior midgut, with Scer\GAL4^Tk.PS

His2Av^HMS02773

adult enteroendocrine cell | decreased number, with Scer\GAL4^Tk.PS

His2Av^HMS02773

anterior crossvein, with Scer\GAL4^dpp.blk1

His2Av^GS3052

cyst cell of testis | somatic clone

His2Av⁸¹⁰

cyst progenitor cell | somatic clone

His2Av⁸¹⁰

egg, with Scer\GAL4^{VP16.mat.αTub67C}

egg chamber, with Scer\GAL4^C587

His2Av^HM05177

egg chamber, with Scer\GAL4^tj.PU

His2Av^HM05177

egg chamber, with Scer\GAL4^{VP16.mat.αTub67C}

eye, with Scer\GAL4^ey.PH

His2Av^HM05177

eye, with Scer\GAL4^GMR.PF

His2Av^HM05177

eye, with Scer\GAL4^hs.2sev

eye | somatic clone

His2Av⁸¹⁰

female germline cyst, with Scer\GAL4^C587

His2Av^HM05177

female germline cyst, with Scer\GAL4^tj.PU

His2Av^HM05177

female germline cyst, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

His2Av^HM05177

germarium, with Scer\GAL4^C587

His2Av^HM05177

germarium, with Scer\GAL4^tj.PU

His2Av^HM05177

germarium | adult stage, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

His2Av^HM05177

imaginal disc

His2Av^KO

lipid droplet | adult stage, with Scer\GAL4^Tk.PS

His2Av^HMS02773

male germline stem cell | somatic clone

His2Av⁸¹⁰

male germline stem cell | somatic clone, with Scer\GAL4^{VP16.nanos.UTR}

His2Av^KK108652

mesonotum, with Scer\GAL4^pnr-MD237

His2Av^GD4747

nurse cell, with Scer\GAL4^{VP16.mat.αTub67C}

ovariole, with Scer\GAL4^C587

His2Av^HM05177

ovariole, with Scer\GAL4^tj.PU

His2Av^HM05177

ovary | adult stage, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

His2Av^HM05177

spermatocyte | somatic clone

His2Av⁸¹⁰

spermatocyte | somatic clone, with Scer\GAL4^{VP16.nanos.UTR}

His2Av^KK108652

tarsal segment, with Scer\GAL4^dpp.blk1

His2Av^GS3127

testis | somatic clone

His2Av⁸¹⁰

testis | somatic clone, with Scer\GAL4^{VP16.nanos.UTR}

His2Av^KK108652

vibrissal bristle, with Scer\GAL4^ey.PH

His2Av^HM05177

wing, with Scer\GAL4^nub.PK

His2Av^HM05177

wing vein, with Scer\GAL4^en.PU

His2Av^HM05177

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (32)

Hsap\H2AZ1

13 of 14

Yes

Hsap\H2AZ2

13 of 14

Yes

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

2 of 14

Yes

Hsap\MACROH2A2

2 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (33)

Rnor\H2az1

13 of 14

Yes

Rnor\H2az2

12 of 14

Yes

Rnor\RGD1562306

7 of 14

Yes

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

Yes

Rnor\Macroh2a2

2 of 14

Yes

Rnor\Rgd1566374

2 of 14

Rnor\Gdap2

1 of 14

Rnor\H2ab3

1 of 14

Yes

Rnor\Hist3h2a

1 of 14

Rnor\LOC685069

1 of 14

Yes

Rnor\LOC689408

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Mus musculus (laboratory mouse) (49)

Mmus\H2az2

13 of 14

Yes

Mmus\H2az1

12 of 14

Yes

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

2 of 14

Yes

Mmus\Macroh2a2

2 of 14

Yes

Mmus\Gdap2

1 of 14

Mmus\H2ab1

1 of 14

Yes

Mmus\H2ab2

1 of 14

Yes

Mmus\H2ab3

1 of 14

Yes

Mmus\H2al1a

1 of 14

Yes

Mmus\H2al1b

1 of 14

Yes

Mmus\H2al1c

1 of 14

Yes

Mmus\H2al1d

1 of 14

Yes

Mmus\H2al1e

1 of 14

Yes

Mmus\H2al1f

1 of 14

Yes

Mmus\H2al1g

1 of 14

Yes

Mmus\H2al1h

1 of 14

Yes

Mmus\H2al1i

1 of 14

Yes

Mmus\H2al1j

1 of 14

Yes

Mmus\H2al1k

1 of 14

Yes

Mmus\H2al1m

1 of 14

Yes

Mmus\H2al1n

1 of 14

Yes

Mmus\H2al1o

1 of 14

Yes

Mmus\H2al2a

1 of 14

Yes

Mmus\H2al2b

1 of 14

Yes

Mmus\H2al2c

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (31)

Xtro\h2az2

10 of 13

Yes

Xtro\h2az1

7 of 13

Yes

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

1 of 13

1 of 13

1 of 13

Yes

Xtro\LOC108648817

1 of 13

Yes

Xtro\LOC108648819

1 of 13

Yes

Xtro\macroh2a2

1 of 13

Yes

Xtro\otud3

1 of 13

Yes

Xtro\ppard

1 of 13

Yes

Danio rerio (Zebrafish) (42)

Drer\h2az2a

13 of 14

Yes

Drer\h2az2b

11 of 14

Yes

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

Drer\si:ch73-368j24.14

3 of 14

Drer\si:dkey-23a13.20

3 of 14

Drer\si:dkey-261m9.11

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

Yes

Drer\si:ch73-368j24.13

2 of 14

Drer\si:ch211-113a14.7

2 of 14

Drer\si:ch211-113a14.11

2 of 14

Drer\si:ch211-113a14.19

2 of 14

Drer\si:ch211-113a14.25

2 of 14

Drer\si:ch1073-153i20.5

2 of 14

Drer\si:ch1073-159d7.10

2 of 14

Drer\si:ch1073-159d7.11

2 of 14

Drer\si:dkey-23a13.6

2 of 14

Drer\si:dkey-108k21.15

2 of 14

Drer\si:dkey-108k21.19

2 of 14

Drer\si:dkey-261m9.7

2 of 14

Drer\si:dkey-261m9.18

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Drer\parp14rs1

1 of 14

Drer\parp14rs4

1 of 14

Yes

Caenorhabditis elegans (Nematode, roundworm) (20)

Cele\htz-1

12 of 14

Yes

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

3 of 14

Yes

Cele\his-57

2 of 14

Cele\B0035.3

1 of 14

Anopheles gambiae (African malaria mosquito) (7)

Agam\AgaP_AGAP000306

12 of 12

Yes

3 of 12

3 of 12

3 of 12

3 of 12

3 of 12

3 of 12

Arabidopsis thaliana (thale-cress) (14)

Atha\HTA8

12 of 13

Yes

Atha\HTA11

12 of 13

Yes

Atha\HTA9

11 of 13

Yes

Atha\HTA4

4 of 13

Yes

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

1 of 13

Saccharomyces cerevisiae (Brewer's yeast) (3)

Scer\HTZ1

12 of 13

Yes

Scer\HTA1

3 of 13

Scer\HTA2

3 of 13

Schizosaccharomyces pombe (Fission yeast) (3)

Spom\pht1

12 of 12

Yes

Spom\hta1

2 of 12

Spom\hta2

2 of 12

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:His2Av. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (20)

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

4 of 13

3 of 13

3 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 ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 1 )

Allele

Disease

Interaction

References

His2Av⁰⁵¹⁴⁶

exacerbates tauopathy

modeled by Hsap\MAPT^GMR.Ex.PJ

(Ambegaokar and Jackson, 2011)

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?

H2AZ1; H2A.Z variant histone 1

13 of 14

142763

H2AZ2; H2A.Z variant histone 2

13 of 14

620008

H2AC1; H2A clustered histone 1

3 of 14

613499

H2AC4; H2A clustered histone 4

3 of 14

602795

H2AC6; H2A clustered histone 6

3 of 14

602794

H2AC7; H2A clustered histone 7

3 of 14

602792

H2AC8; H2A clustered histone 8

3 of 14

602786

H2AC11; H2A clustered histone 11

3 of 14

615012

H2AC12; H2A clustered histone 12

3 of 14

615013

H2AC13; H2A clustered histone 13

3 of 14

602787

H2AC14; H2A clustered histone 14

3 of 14

602791

H2AC15; H2A clustered histone 15

3 of 14

602788

H2AC16; H2A clustered histone 16

3 of 14

602793

H2AC17; H2A clustered histone 17

3 of 14

602796

H2AC18; H2A clustered histone 18

3 of 14

142720

H2AC19; H2A clustered histone 19

3 of 14

H2AC20; H2A clustered histone 20

3 of 14

602797

H2AC21; H2A clustered histone 21

3 of 14

615014

H2AJ; H2A.J histone

3 of 14

H2AC25; H2A clustered histone 25

3 of 14

615015

H2AX; H2A.X variant histone

3 of 14

601772

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

View in FlyBase Interactions Browser View in MIST tool (external link)

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

His2Av

split luciferase complementation, luminiscence technology

(Kolkhof et al., 2017)

His2Av - CG11700

split luciferase complementation, luminiscence technology

(Kolkhof et al., 2017)

His2Av - CG8677

anti bait coimmunoprecipitation, western blot

(Hanai et al., 2008)

His2Av - His2A

split luciferase complementation, luminiscence technology

(Kolkhof et al., 2017)

His2Av - His2B

split luciferase complementation, luminiscence technology

(Kolkhof et al., 2017)

His2Av - His4

split luciferase complementation, luminiscence technology

(Kolkhof et al., 2017)

His2Av - Imp

pull down, western blot

(Dronamraju and Mason, 2009)

His2Av - JIL-1

pull down, western blot, enzymatic study, autoradiography

(Thomas et al., 2014)

His2Av - Jabba

split luciferase complementation, luminiscence technology, anti tag coimmunoprecipitation, western blot

(Stephenson et al., 2021, Kolkhof et al., 2017, Li et al., 2012)

His2Av - Parp1

anti tag coimmunoprecipitation, western blot

(Thomas et al., 2019)

His2Av - Pc

affinity technology, peptide array, western blot

(Stein et al., 2013)

His2Av - Su(var)2-10

anti tag coimmunoprecipitation, anti tag western blot

(Tang et al., 2021)

His2Av - Su(var)205

anti tag coimmunoprecipitation, western blot, pull down, anti tag western blot

(Vernì and Cenci, 2015)

His2Av - Tip60

enzymatic study, western blot, anti bait coimmunoprecipitation

(Kusch et al., 2014, Kusch et al., 2004)

His2Av - Yeti

anti tag coimmunoprecipitation, western blot

(Messina et al., 2014)

His2Av - mu2

pull down, molecular weight estimation by coomasie staining, Identification by mass spectrometry

(Dronamraju and Mason, 2009)

His2Av - nbs

pull down, western blot

(Dronamraju and Mason, 2009)

His2Av - pont

anti tag coimmunoprecipitation, western blot, peptide massfingerprinting

(Kusch et al., 2004)

His2Av - rad50

pull down, western blot

(Dronamraju and Mason, 2009)

His2Av - tefu

enzymatic study, autoradiography, anti tag coimmunoprecipitation, anti tag western blot

(Hong and Choi, 2013)

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

His2Av

enhanceable

CG8677

(Hanai et al., 2008)

His2Av

enhanceable

dom

(Morillo Prado et al., 2013)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

aurA

suppressible

His2Av

(Merigliano et al., 2019)

CG8677

enhanceable

His2Av

(Hanai et al., 2008)

enhanceable

(Ji et al., 2012)

suppressible

(Ji et al., 2012)

enhanceable

(Ji et al., 2012)

enhanceable

(Gregory et al., 2007)

Ras85D

suppressible

His2Av

(Murcia et al., 2019)

trx|ash1

suppressible

His2Av

(Swaminathan et al., 2005)

upd1

suppressible

His2Av

(Bach et al., 2003)

External Data

Subunit Structure (UniProtKB)

(UniProt, P08985 )

Linkouts

BioGRID - A database of protein and genetic interactions.

68120

DroID - A comprehensive database of gene and protein interactions.

FBgn0001197

MIST (genetic) - An integrated Molecular Interaction Database

43229

MIST (protein-protein) - An integrated Molecular Interaction Database

43229

Pathways

Signaling Pathways (FlyBase)

NEGATIVE REGULATORS OF IMD SIGNALING PATHWAY

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.

ATP-dependent chromatin remodeling

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

3-92

Cytogenetic map

97D3-97D3

Sequence location

3R:26,866,929..26,869,220 [+]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

97D3-97D3

Limits computationally determined from genome sequence between P{lacW}scrib^j7B3 and P{lacW}His2Av^L1602

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

97D3-97D6

(determined by in situ hybridisation)

(Spradling et al., 1999)

97D3-97D6

(BDGP Project Members, 1994-1999)

97C-97D

(determined by in situ hybridisation)

(van Daal et al., 1988)

Experimentally Determined Recombination Data

Location

3-92

(FlyBase, 2016)

3-88.9

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

The order from centromere to telomere is l(3)97Da l(3)97Db l(3)97Dc. His2Av is likely to map near to l(3)97Db.

(Knibb et al., 1993)

Stocks and Reagents

Stocks (42)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

53731

w^*; P{UAS-His2Av.mKO}3

23651

w^*; P{His2Av-mRFP1}II.2

91659

w^*; P{His2Av-mCer}III.1

94935

w^*; P{His2Av-iRFP713}L5

94936

w^*; P{His2Av-iRFP713}L2

23650

w^*; P{His2Av-mRFP1}III.1

94934

P{His2Av-2xiRFP713}1, w^*

606943

w^*; P{His2Av.Halo}attP40

28966

y¹ v¹; P{TRiP.HM05177}attP2

Kyoto

108494

w¹¹¹⁸; P{His2Av.GFP(S65T)}62A

VDRC

v110598

P{KK108652}VIE-260B

v12768

w¹¹¹⁸; P{GD4747}v12768

More stocks available...

Genomic Clones (14)

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

cDNA Clones (156)

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

LD21568

Other clones

DPiM_LD21568

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.

FBgn0001197

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0001197

Antibody Information

Laboratory Generated Antibodies

monoclonal

(González-Marín et al., 2023, Lake et al., 2013)

polyclonal

(Börner and Becker, 2016, Jeon et al., 2015, Lancaster et al., 2010, Mehrotra and McKim, 2006, Madigan et al., 2002, van Daal and Elgin, 1992)

Commercially Available Antibodies

Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research

UNC93-5.2.1

Cell Line Information

Publicly Available Cell Lines

S2R+-His2Av-mNeonGreen-B11

Other Stable Cell Lines

Other Comments

His2Av is sufficient for Parp protein localisation.

(Kotova et al., 2011)

Activation of Parp in vivo and the start of Parp-dependent processes in chromatin are triggered by phosphorylation of His2Av.

(Kotova et al., 2011)

Phosphorylation of His2Av is critical for fly viability after gamma-irradiation.

(Kotova et al., 2011)

The C terminal region of His2Av, excluding the last 12 amino acids, is critical for function. This region is buried inside the histone core and is not responsible for interactions with DNA, but is possibly involved in protein-protein interactions.

(Clarkson et al., 1999)

In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes.

(Kliman and Eyre-Walker, 1998)

His2Av function is essential for nucleosome assembly and is continuously expressed.

(van Daal and Elgin, 1992)

His2Av has been cloned and sequenced.

(van Daal et al., 1990)

Isolated from a cDNA library, using a Tetrahymena histone H2A variant hv1 cDNA as a probe.

(van Daal et al., 1988)

A His2Av cDNA has been cloned and sequenced.

(van Daal et al., 1988)

Relationship to Other Genes

Source for database merge of

Source for merge of: His2Av l(3)05146

(Spradling et al., 1999)

Additional comments

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (68)

Reported As

Symbol Synonym

*i H2av

(Duy Binh et al., 2019)

5499

(Ravi et al., 2009)

CG5499

(Rosa-Ferreira et al., 2015, Jaspers et al., 2012, Cammarato et al., 2011, Kühnlein, 2011, Ni et al., 2011.7.19, Müller et al., 2010, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009, Quinones-Coello, 2007, Quinones-Coello, 2007, Wagner et al., 2007, Cermelli et al., 2006, Laviolette et al., 2005, Kusch et al., 2004, Rebollo et al., 2004)

H2A

(Feng et al., 2014, Wang et al., 2011, Schaefer et al., 2008)

H2A.F/Z

(Zhu et al., 2007)

H2A.V

(Scacchetti et al., 2020, Harpprecht et al., 2019, Khuong et al., 2017, Börner and Becker, 2016, Cipressa et al., 2016, Messina et al., 2016, Vernì and Cenci, 2015, Messina et al., 2014, Ida et al., 2013)

H2A.X

H2A.Z

(Janssens et al., 2017, Yang and Ioshikhes, 2017, Derive et al., 2015, Horard and Loppin, 2015, Schauer et al., 2013, Xing et al., 2013, Egelhofer et al., 2011, Rach et al., 2011, Weber et al., 2010, Cui and Zhurkin, 2009, Frydrychova et al., 2008, Mavrich et al., 2008, Raisner and Madhani, 2006, Varga-Weisz and Becker, 2006, Lippman and Martienssen, 2004)

H2A.v

(Santopietro et al., 2025)

H2AV

(Becker, 2024, Jayakrishnan et al., 2024, Cicconi et al., 2017, Lake et al., 2013, Stein et al., 2013, Nagel et al., 2012, Sakai et al., 2009, Klovstad et al., 2008, Mehrotra et al., 2008, Garcia et al., 2007, Bandura, 2005, Purdy et al., 2005)

H2AV_DROM

(Storm and Sonnhammer, 2003)

H2AX

(de la Cova et al., 2014, Yan et al., 2011, Klattenhoff et al., 2009, Andreyeva et al., 2008, Klattenhoff et al., 2007)

H2AZ

(Redon et al., 2002)

H2Av

(Balachandra and Amodeo, 2025, Chen et al., 2025, Shindo et al., 2025, Vicars et al., 2025, Bamgbose and Tulin, 2024, Zhimulev et al., 2024, Bernasek et al., 2023, Corcoran and Jacob, 2023, Dubruille et al., 2023, Mari et al., 2023, Tirgar et al., 2023, Warecki et al., 2023, Zhang et al., 2023, Bernués et al., 2022, Cammarata-Mouchtouris et al., 2022, Chakravarti et al., 2022, de-Carvalho et al., 2022, Feng et al., 2022, Molano-Fernández et al., 2022, Sang et al., 2022, Shindo et al., 2022, Simmons et al., 2022, Xu et al., 2022, Llorens-Giralt et al., 2021, Pitchakarn et al., 2021, Saunders, 2021, Stephenson et al., 2021, Tang et al., 2021, Vicars et al., 2021, Vigano et al., 2021, Meyer-Nava et al., 2020, Miwa et al., 2020, Palladino et al., 2020, Warecki et al., 2020, Billmyre et al., 2019, Merigliano et al., 2019, Mirkovic et al., 2019, Mota et al., 2019, Park et al., 2019, Thomas et al., 2019, Harumoto and Lemaitre, 2018, Johnson et al., 2018, Rust et al., 2018, Yao et al., 2018, Karam et al., 2017, Kolkhof et al., 2017, Li et al., 2017, Yang et al., 2017, Müller, 2016.10.7, Ryu et al., 2016, Alexander et al., 2015, Dietz et al., 2015, Horard and Loppin, 2015, Marin-Vicente et al., 2015, Pengelly et al., 2015, Amcheslavsky et al., 2014, Frost et al., 2014, Ho et al., 2014, Kuang et al., 2014, Landais et al., 2014, Rathke et al., 2014, Thomas et al., 2014, Fujita et al., 2013, Jagut et al., 2013, Kimura, 2013, Pérez-Montero et al., 2013, Petersen et al., 2013, Silver-Morse and Li, 2013, Agelopoulos et al., 2012, Althoff et al., 2012, Chmielewski et al., 2012, Ferguson et al., 2012, Li et al., 2012, Li et al., 2012, Muha et al., 2012, Pek et al., 2012, Petersen et al., 2012, Vujatovic et al., 2012, Kanesaki et al., 2011, Kotova et al., 2011, Kriston-Vizi et al., 2011, Redhouse et al., 2011, Yan et al., 2011, Zhang and Pugh, 2011, Zhang et al., 2011, Bryson et al., 2010, Chioda et al., 2010, Deal et al., 2010, Huang et al., 2010, Macalpine et al., 2010, Portela et al., 2010, Schittenhelm et al., 2010, Weber et al., 2010, Andreyenkova et al., 2009, Clapier and Cairns, 2009, Dronamraju and Mason, 2009, Henikoff et al., 2009, Klapholz et al., 2009, McCleland et al., 2009, Narbonne-Reveau and Lilly, 2009, Nie et al., 2009, Peng and Karpen, 2009, Andreyeva et al., 2008, Hanai et al., 2008, Langley and Karpen, 2008, Martinek et al., 2008, Mavrich et al., 2008, Pauli et al., 2008, Royou et al., 2008, Zhang and Karpen, 2008, Haynes et al., 2007, Hong et al., 2007, Johansson et al., 2007, Klattenhoff et al., 2007, Langley and Karpen, 2007, Lu et al., 2007, Oikemus et al., 2007, Pandey et al., 2007, Pinnola et al., 2007, Sakaguchi and Steward, 2007, Takada et al., 2007, Tulin et al., 2007, Wagner et al., 2007, Wagner et al., 2007, Bouazoune and Brehm, 2006, Brasaemle and Hansen, 2006, Cermelli et al., 2006, Polo and Almouzni, 2006, Heeger et al., 2005, Ivanovska et al., 2005, Jin, 2005, Kusch et al., 2004, Kusch et al., 2004, Kusch et al., 2004, Webber et al., 2004, Glaser et al., 2002, Madigan et al., 2002, Leach et al., 2000)

H2a.V

(Wagner et al., 2007)

H2av

(Lam et al., 2005)

H2avD

(Wen et al., 2008, Manak et al., 2007)

HIS

(Di Talia and Wieschaus, 2012, Di Talia and Wieschaus, 2012)

HIS2AV

(Riddle et al., 2012, Joyce and McKim, 2009, Joyce et al., 2009, McCaffrey et al., 2006)

HIS2AVD

(Staudt et al., 2006)

His

(Althoff et al., 2012)

His2

(Ninov et al., 2010)

His2A

(Foronda et al., 2012, Schaefer et al., 2008)

His2AV

(Villa-Fombuena et al., 2021, Salvany et al., 2012, Blumenstiel et al., 2009)

His2Av

(Lee et al., 2025, Marsh et al., 2025, Molano-Fernández et al., 2025, Neophytou et al., 2025, Stockinger et al., 2025, Tang and Wu, 2025.8.22, Xu et al., 2025, Bellec et al., 2024, Choudhury et al., 2024, Hiraiwa et al., 2024, Perlegos et al., 2024, Forbes Beadle et al., 2023, Berry et al., 2022, Cho et al., 2022, Falo-Sanjuan and Bray, 2022, Ferree et al., 2022, Kabakci et al., 2022, Kakanj et al., 2022, Logeay et al., 2022, Marchetti et al., 2022, Pan et al., 2022, Yuen et al., 2022, Zong et al., 2022, Fukaya, 2021, Reilein et al., 2021, Sterne et al., 2021, Wilcockson and Ashe, 2021, Witt et al., 2021, Bosch et al., 2020, Pirani et al., 2020, Weber et al., 2020, Bouska et al., 2019, Deneke et al., 2019, Kondo and Hayashi, 2019, McDaniel et al., 2019, Murcia et al., 2019, Wei et al., 2019, Witt et al., 2019, Cruz-Becerra et al., 2018, Giaimo et al., 2018, Lim et al., 2018, Arata et al., 2017, Cheng et al., 2017, Fukaya et al., 2017, Blattner et al., 2016, Hampoelz et al., 2016, Ozawa et al., 2016, Tatomer et al., 2016, Blythe and Wieschaus, 2015, Dorn and Dorn, 2015, Kwong et al., 2015, Hughes et al., 2014, Radermacher et al., 2014, Radermacher et al., 2014, Czech et al., 2013, Debruhl et al., 2013, Di Talia et al., 2013, Di Talia et al., 2013, Handler et al., 2013, Herzog et al., 2013, Hong and Choi, 2013, Kockmann et al., 2013, Kwon et al., 2013, Morillo Prado et al., 2013, Sakurai et al., 2013, Althoff et al., 2012, Comeron et al., 2012, Di Talia and Wieschaus, 2012, Fasulo et al., 2012, Foronda et al., 2012, Kanakousaki and Gibson, 2012, Telley et al., 2012, Abreu-Blanco et al., 2011, Ambegaokar and Jackson, 2011, Friedman et al., 2011, Joyce et al., 2011, Kotova et al., 2011, Kuranaga et al., 2011, Lake et al., 2011, Little et al., 2011, Lott et al., 2011, Morris and Spradling, 2011, Taliaferro et al., 2011, Kong et al., 2010, Müller et al., 2010, Baldinger and Gossen, 2009, Bischoff and Cseresnyés, 2009, McCleland et al., 2009, Nakayama et al., 2009, Sing et al., 2009, Venken et al., 2009, Christensen et al., 2008.4.15, Christensen et al., 2008.6.11, Hanai et al., 2008, Maurange et al., 2008, McElroy et al., 2008, Pauli et al., 2008, Schaefer et al., 2008, Titen and Golic, 2008, Buszczak et al., 2007, Gregory et al., 2007, Heidmann, 2007.2, Lake et al., 2007, Langille and Clark, 2007, Quinones-Coello, 2007, Schubiger, 2007.9.19, Schuh et al., 2007, Schuh et al., 2007, Zhong and Belote, 2007, Beller et al., 2006, Dorus et al., 2006, Dudu, 2006, Dudu et al., 2006, Mehrotra and McKim, 2006, Laviolette et al., 2005, Raja and Renkawitz-Pohl, 2005, Jang et al., 2003)

His2aV

(Tatomer et al., 2016)

His2av

(Dufourt et al., 2021, Merigliano et al., 2017)

HisH2Av

(Andreyeva et al., 2008, Oliveira et al., 2007, Quinones-Coello, 2007)

Hist

(Tipping et al., 2010, Coppey et al., 2008)

Hist2av

(Parma et al., 2007)

P-H2Av

(Joy et al., 2021)

dHis2Av

(Goodman et al., 2019)

ggr;-H2Av

(Klemm et al., 2025)

h2AvD

(Gao et al., 2009, Rong, 2008)

his

(Pokrywka et al., 2009)

his2Av

(Smibert et al., 2012)

his2av

(Angulo et al., 2019)

l(3)05146

(Kidd et al., 1999, Meister and Braun, 1995.10, BDGP Project Members, 1994-1999, Berkeley Drosophila Genome Project, 1993.6.1)

l(3)810

l(3)97Dd

(Yasuda et al., 1995, Knibb et al., 1993)

l(3)L1602

(Beaton, 2000.1.31)

γ-H2AV

(Kotov et al., 2024, Lake et al., 2015)

γ-H2Av

(Nguyen et al., 2024, Ryu et al., 2024, Watase and Yamashita, 2024, Wang et al., 2021, Mascolo et al., 2020, Oberacker et al., 2018, Yeom et al., 2015, Zhang et al., 2015, Loh et al., 2012, Klattenhoff et al., 2009, Park and Asano, 2008)

γ-H2AvD

(Karling and Weavers, 2025, Park et al., 2024, Na et al., 2018)

γ-HIS2AV

(Gong et al., 2005)

γ-His2Av

(Jagut et al., 2013, Peretz et al., 2009, Mehrotra and McKim, 2006)

γH2AV

(Apostolou et al., 2025, Lake et al., 2015)

γH2Av

(Nirala et al., 2021, Bhargava et al., 2020, Park and Kim, 2020, Amaral et al., 2017, Khan et al., 2017, Sengupta et al., 2016, Ryu et al., 2015, Jagut et al., 2013, Lancaster et al., 2010)

γH2Ax

(Khan et al., 2017)

γH2av

(Nagy and Buchon, 2025)

γH2ax

(Khan et al., 2019)

Name Synonyms

H2A variant

(Kaeser and Emerson, 2006)

His2Av

(Marie et al., 2017)

Histone

(Zheng et al., 2013)

Histone H2A variant

(Cammarato et al., 2011, Kühnlein, 2011)

Histone H2A.v

(Urban et al., 2017)

Histone-2A

(Taliaferro et al., 2011)

Histone-2av

(Chinta et al., 2012)

Histone2Av

(Schmidt et al., 2021, Tsui et al., 2016, Kuranaga et al., 2011)

histone

(Wang et al., 2013, Puah et al., 2011, Wang et al., 2010, McCleland et al., 2009, Coppey et al., 2008, McCleland and O'Farrell, 2008, Frydman et al., 2006, Yu et al., 2006, Savoian and Rieder, 2002)

histone 2A variant

(Lake et al., 2015, Schuh et al., 2007)

histone H2A variant

(Chioda et al., 2010, Pandey et al., 2007, Cermelli et al., 2006)

histone H2Av

(Kondo and Perrimon, 2011, Pandey et al., 2007)

histone variant H2Av

(Bulchand et al., 2010)

histone2A

(Skold et al., 2005)

histone2Av

(Weislogel et al., 2013)

histone2AvD

(Schneider et al., 2006)

Secondary FlyBase IDs

FBgn0010859

Datasets (2)

Study focus (2)

Experimental Role

Project

Project Type

Title

RNAi_target

modENCODE_chromatin_histones_cells

genome binding

Genome-wide localization of histones and their modifications in cell lines by ChIP-chip and ChIP-Seq.

bait_protein

modENCODE_chromatin_histones_flies

genome binding

Genome-wide localization of histones and their modifications in fly tissues by ChIP-chip and ChIP-Seq.

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 67 )

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.

43229

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.

P08985

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

P08985

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.

P08985

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0001197

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

43229/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0001197

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0001197

FlyMine - An integrated database for Drosophila genomics

FBgn0001197

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG5499

MARRVEL_MODEL - MARRVEL (model organism gene)

43229

Linkouts

BioGRID - A database of protein and genetic interactions.

68120

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0001197

DroID - A comprehensive database of gene and protein interactions.

FBgn0001197

DRSC - Results frm RNAi screens

FBgn0001197

Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research

UNC93-5.2.1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0001197

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0001197

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG5499

Flygut - An atlas of the Drosophila adult midgut

FBgn0001197

FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.

FBgn0001197

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC000931

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.

ATP-dependent chromatin remodeling

MIST (genetic) - An integrated Molecular Interaction Database

43229

MIST (protein-protein) - An integrated Molecular Interaction Database

43229

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

References (642)

The following external sites may use different assemblies or annotations than FlyBase.

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