Gene Symbol: His2Av
Description: Histone H2A variant
Alias: 5499, CG5499, Dmel\CG5499, H2A, H2A.F/Z, H2A.V, H2A.X, H2A.Z, H2AV, H2AV_DROM, H2AX, H2AZ, H2Av, H2AvD, H2a.V, H2av, H2avD, HIS, HIS2AV, HIS2AVD, His, His2, His2A, His2AV, His2AvD, HisH2Av, Hist, Hist2av, gamma-H2AV, gamma-H2Av, gamma-HIS2AV, gamma-His2Av, gammaH2AV, gammaH2Av, h2AvD, his, his2Av, l(3)05146, l(3)810, l(3)97Dd, l(3)L1602, histone H2A , CG5499-PA, CG5499-PB, H2A , His2Av-PA, His2Av-PB, Histone-2A, Histone-2av, histone, histone , histone 2A , histone H2Av, histone2A, histone2Av, histone2AvD
Species: fruit fly
Products:     His2Av

Top Publications

  1. Clarkson M, Wells J, Gibson F, Saint R, Tremethick D. Regions of variant histone His2AvD required for Drosophila development. Nature. 1999;399:694-7 pubmed
    ..domain could be established to carry out a specialized function is by the localized incorporation of specific histone variants into nucleosomes...
  2. Wasser M, Chia W. The Drosophila EAST protein associates with a nuclear remnant during mitosis and constrains chromosome mobility. J Cell Sci. 2003;116:1733-43 pubmed
    ..We propose that EAST constitutes a component of a nucleoskeleton that helps to constrain the mobility of chromosomes in interphase, mitosis and meiosis. ..
  3. McCleland M, Shermoen A, O Farrell P. DNA replication times the cell cycle and contributes to the mid-blastula transition in Drosophila embryos. J Cell Biol. 2009;187:7-14 pubmed publisher
    ..We conclude that normal S phase and replication checkpoint activities are important timers of the undisturbed cell cycle before, but not after, the MBT...
  4. Katsani K, Karess R, Dostatni N, Doye V. In vivo dynamics of Drosophila nuclear envelope components. Mol Biol Cell. 2008;19:3652-66 pubmed publisher
    ..Surprisingly, Drosophila Nup107 shows no evidence of localization to kinetochores, despite the demonstrated importance of this association in mammalian cells. ..
  5. Corona D, Siriaco G, Armstrong J, Snarskaya N, McClymont S, Scott M, et al. ISWI regulates higher-order chromatin structure and histone H1 assembly in vivo. PLoS Biol. 2007;5:e232 pubmed
    ..does not cause obvious defects in nucleosome assembly, but results in a significant reduction in the level of histone H1 associated with chromatin in vivo...
  6. Royou A, McCusker D, Kellogg D, Sullivan W. Grapes(Chk1) prevents nuclear CDK1 activation by delaying cyclin B nuclear accumulation. J Cell Biol. 2008;183:63-75 pubmed publisher
    ..Grp(Chk1) prevents nuclear CDK1 activation by delaying CycB nuclear accumulation through Wee1-dependent and independent mechanisms. ..
  7. Henikoff S, Henikoff J, Sakai A, Loeb G, Ahmad K. Genome-wide profiling of salt fractions maps physical properties of chromatin. Genome Res. 2009;19:460-9 pubmed publisher
    ..3 and correspond closely to profiles of histone H2Av (H2A.Z) and RNA polymerase II. This correspondence suggests that transcription can result in loss of H3...
  8. Narbonne Reveau K, Lilly M. The Cyclin-dependent kinase inhibitor Dacapo promotes genomic stability during premeiotic S phase. Mol Biol Cell. 2009;20:1960-9 pubmed publisher
    ..Finally, we report that dap(-/-) ovarian cysts frequently undergo an extramitotic division before meiotic entry, indicating that Dap influences the timing of the mitotic/meiotic transition. ..
  9. Savoian M, Rieder C. Mitosis in primary cultures of Drosophila melanogaster larval neuroblasts. J Cell Sci. 2002;115:3061-72 pubmed
    ..5 minutes. Analysis of neuroblasts containing various GFP-tagged proteins (e.g. histone, fizzy, fizzy-related and alpha-tubulin) reveals that attaching kinetochores exhibit sudden, rapid pole-directed ..

More Information


  1. Chinta R, Wasser M. Three-dimensional segmentation of nuclei and mitotic chromosomes for the study of cell divisions in live Drosophila embryos. Cytometry A. 2012;81:52-64 pubmed publisher
    ..We tested our method on 3D time-series images of live embryos expressing histone-2Av-green fluorescence protein...
  2. Klattenhoff C, Bratu D, McGinnis Schultz N, Koppetsch B, Cook H, Theurkauf W. Drosophila rasiRNA pathway mutations disrupt embryonic axis specification through activation of an ATR/Chk2 DNA damage response. Dev Cell. 2007;12:45-55 pubmed
    ..Furthermore, rasiRNA pathway mutations lead to germline-specific accumulation of gamma-H2Av foci characteristic of DNA damage...
  3. Swaminathan J, Baxter E, Corces V. The role of histone H2Av variant replacement and histone H4 acetylation in the establishment of Drosophila heterochromatin. Genes Dev. 2005;19:65-76 pubmed
    ..Here we show that the histone H2A variant of Drosophila melanogaster, H2Av, localizes to the centromeric heterochromatin, and it is recruited to ..
  4. Parry D, O Farrell P. The schedule of destruction of three mitotic cyclins can dictate the timing of events during exit from mitosis. Curr Biol. 2001;11:671-83 pubmed
    ..We suggest that the schedule of degradation of cyclin A, cyclin B, and then cyclin B3 contributes to the temporal coordination of mitotic events. ..
  5. Siriaco G, Deuring R, Chioda M, Becker P, Tamkun J. Drosophila ISWI regulates the association of histone H1 with interphase chromosomes in vivo. Genetics. 2009;182:661-9 pubmed publisher
    ..chromatin-remodeling factor plays a key role in this process by promoting the assembly of chromatin containing histone H1. To test this hypothesis, we investigated the function of H1 in Drosophila...
  6. Jayaramaiah Raja S, Renkawitz Pohl R. Replacement by Drosophila melanogaster protamines and Mst77F of histones during chromatin condensation in late spermatids and role of sesame in the removal of these proteins from the male pronucleus. Mol Cell Biol. 2005;25:6165-77 pubmed
    ..Mst77F encodes a linker histone-like protein showing significant similarity to mammalian HILS1 protein...
  7. Joyce E, Pedersen M, Tiong S, White Brown S, Paul A, Campbell S, et al. Drosophila ATM and ATR have distinct activities in the regulation of meiotic DNA damage and repair. J Cell Biol. 2011;195:359-67 pubmed publisher
    ..Both kinases phosphorylate H2AV (?-H2AV), and, using this as a reporter for ATM/ATR activity, we found that the DSB repair response is surprisingly ..
  8. Schittenhelm R, Althoff F, Heidmann S, Lehner C. Detrimental incorporation of excess Cenp-A/Cid and Cenp-C into Drosophila centromeres is prevented by limiting amounts of the bridging factor Cal1. J Cell Sci. 2010;123:3768-79 pubmed publisher
    ..cycle progression in higher eukaryotes depends critically on the faithful incorporation of a centromere-specific histone H3 variant encoded by CENPA in humans and cid in Drosophila...
  9. Pauli A, Althoff F, Oliveira R, Heidmann S, Schuldiner O, Lehner C, et al. Cell-type-specific TEV protease cleavage reveals cohesin functions in Drosophila neurons. Dev Cell. 2008;14:239-51 pubmed publisher
    ..These data demonstrate essential roles for cohesin in nondividing cells and also introduce a powerful tool by which to investigate protein function in metazoa. ..
  10. Heeger S, Leismann O, Schittenhelm R, Schraidt O, Heidmann S, Lehner C. Genetic interactions of separase regulatory subunits reveal the diverged Drosophila Cenp-C homolog. Genes Dev. 2005;19:2041-53 pubmed
  11. Puah W, Cheok L, Biro M, Ng W, Wasser M. TLM-Converter: reorganization of long time-lapse microscopy datasets for downstream image analysis. Biotechniques. 2011;51:49-50, 52-3 pubmed publisher
    ..In addition, saving the outputs in open standard image file formats enables data sharing between independently developed software tools. ..
  12. Madigan J, Chotkowski H, Glaser R. DNA double-strand break-induced phosphorylation of Drosophila histone variant H2Av helps prevent radiation-induced apoptosis. Nucleic Acids Res. 2002;30:3698-705 pubmed
    ..Histone variant H2AX and core histone H2A are phosphorylated in mammals and budding yeast, respectively...
  13. Pandey R, Heeger S, Lehner C. Rapid effects of acute anoxia on spindle kinetochore interactions activate the mitotic spindle checkpoint. J Cell Sci. 2007;120:2807-18 pubmed
    ..Although involving adenosine triphosphate (ATP)-consuming kinases, the spindle checkpoint can therefore be activated by spindle damage in response to acute anoxia and protect against aneuploidies. ..
  14. Schuh M, Lehner C, Heidmann S. Incorporation of Drosophila CID/CENP-A and CENP-C into centromeres during early embryonic anaphase. Curr Biol. 2007;17:237-43 pubmed
    ..A candidate for an epigenetic mark is the centromere-specific histone H3 variant CENP-A that replaces H3 in alternating blocks of chromatin exclusively in active centromeres...
  15. Ji J, Squirrell J, Schubiger G. Both cyclin B levels and DNA-replication checkpoint control the early embryonic mitoses in Drosophila. Development. 2004;131:401-11 pubmed
  16. Crest J, Oxnard N, Ji J, Schubiger G. Onset of the DNA replication checkpoint in the early Drosophila embryo. Genetics. 2007;175:567-84 pubmed
    ..Our results suggest an antagonistic interaction between DNA replication checkpoint activation and Cdk1-CycB activity during the transition from preblastoderm to blastoderm cycles. ..
  17. Wang H, Brust Mascher I, Cheerambathur D, Scholey J. Coupling between microtubule sliding, plus-end growth and spindle length revealed by kinesin-8 depletion. Cytoskeleton (Hoboken). 2010;67:715-28 pubmed publisher
  18. Joyce E, McKim K. Drosophila PCH2 is required for a pachytene checkpoint that monitors double-strand-break-independent events leading to meiotic crossover formation. Genetics. 2009;181:39-51 pubmed publisher
    ..Interestingly, PCH2-dependent delays in prophase may allow additional crossovers to form. ..
  19. Telley I, Gaspar I, Ephrussi A, Surrey T. Aster migration determines the length scale of nuclear separation in the Drosophila syncytial embryo. J Cell Biol. 2012;197:887-95 pubmed publisher
    ..These activities define a characteristic separation length scale that appears to be a conserved property of developing insect embryos. ..
  20. Fyodorov D, Blower M, Karpen G, Kadonaga J. Acf1 confers unique activities to ACF/CHRAC and promotes the formation rather than disruption of chromatin in vivo. Genes Dev. 2004;18:170-83 pubmed
    ..These findings collectively indicate that ACF/CHRAC functions in the assembly of periodic nucleosome arrays that contribute to the repression of genetic activity in the eukaryotic nucleus. ..
  21. Yu J, Fleming S, Williams B, Williams E, Li Z, Somma P, et al. Greatwall kinase: a nuclear protein required for proper chromosome condensation and mitotic progression in Drosophila. J Cell Biol. 2004;164:487-92 pubmed
    ..Chromosomes are highly undercondensed, particularly in the euchromatin, but nevertheless contain phosphorylated histone H3, condensin, and topoisomerase II...
  22. Purdy A, Uyetake L, Cordeiro M, Su T. Regulation of mitosis in response to damaged or incompletely replicated DNA require different levels of Grapes (Drosophila Chk1). J Cell Sci. 2005;118:3305-15 pubmed
    ..Finally, our results demonstrate that Drosophila provides a unique opportunity to study the regulation of the entry into, and progress through, mitosis by DNA structure checkpoints in metazoa. ..
  23. Varga Weisz P, Becker P. Regulation of higher-order chromatin structures by nucleosome-remodelling factors. Curr Opin Genet Dev. 2006;16:151-6 pubmed
    ..At the level of single nucleosomes, they are involved in nucleosome-repositioning, altering histone-DNA interactions, disassembly of nucleosomes, and the exchange of histones with variants of different properties...
  24. Lake C, Nielsen R, Hawley R. The Drosophila zinc finger protein trade embargo is required for double strand break formation in meiosis. PLoS Genet. 2011;7:e1002005 pubmed publisher
    ..We speculate that Trem may act by either directing the binding of Mei-P22 to preferred sites of DSB formation or by altering chromatin structure in a manner that allows Mei-P22 to form foci. ..
  25. McCleland M, Farrell J, O Farrell P. Influence of cyclin type and dose on mitotic entry and progression in the early Drosophila embryo. J Cell Biol. 2009;184:639-46 pubmed publisher
  26. Mehrotra S, Maqbool S, Kolpakas A, Murnen K, Calvi B. Endocycling cells do not apoptose in response to DNA rereplication genotoxic stress. Genes Dev. 2008;22:3158-71 pubmed publisher
    ..Similar mechanisms may operate during vertebrate development, with implications for cancer predisposition in certain tissues. ..
  27. Fischer M, Heeger S, Hacker U, Lehner C. The mitotic arrest in response to hypoxia and of polar bodies during early embryogenesis requires Drosophila Mps1. Curr Biol. 2004;14:2019-24 pubmed
    ..Finally, we show that Mps1 and the mitotic spindle checkpoint are responsible for the developmental cell cycle arrest of the three haploid products of female meiosis that are not used as the female pronucleus. ..
  28. Polo S, Almouzni G. Chromatin assembly: a basic recipe with various flavours. Curr Opin Genet Dev. 2006;16:104-11 pubmed
    Packaging of eukaryotic genomes into chromatin is a hierarchical mechanism, starting with histone deposition onto DNA to produce nucleosome arrays, which then further fold and ultimately form functional domains...
  29. Echard A, O Farrell P. The degradation of two mitotic cyclins contributes to the timing of cytokinesis. Curr Biol. 2003;13:373-83 pubmed
    ..Stable cyclin B3 also blocks the later return to interphase that otherwise appears to impose a deadline for the completion of cytokinesis. ..
  30. Jang J, Sherizen D, Bhagat R, Manheim E, McKim K. Relationship of DNA double-strand breaks to synapsis in Drosophila. J Cell Sci. 2003;116:3069-77 pubmed
    ..requirements for double-strand break formation during Drosophila meiosis, we used an antibody that recognizes a histone modification at double-strand break sites, phosphorylation of HIS2AV (gamma-HIS2AV)...
  31. Webber H, Howard L, Bickel S. The cohesion protein ORD is required for homologue bias during meiotic recombination. J Cell Biol. 2004;164:819-29 pubmed
    ..We conclude that ORD activity suppresses sister chromatid exchange and stimulates inter-homologue crossovers, thereby promoting homologue bias during meiotic recombination in Drosophila. ..
  32. Oliveira R, Heidmann S, Sunkel C. Condensin I binds chromatin early in prophase and displays a highly dynamic association with Drosophila mitotic chromosomes. Chromosoma. 2007;116:259-74 pubmed
    ..Furthermore, the rapid turnover of Barren-EGFP indicates that the mechanism by which condensin I promotes mitotic chromosome organization is inconsistent with a static scaffold model. ..
  33. Wasser M, Chia W. The EAST protein of drosophila controls an expandable nuclear endoskeleton. Nat Cell Biol. 2000;2:268-75 pubmed
    ..We propose that EAST mediates the assembly of an expandable nuclear endoskeleton which, through alterations of its volume, can modulate the spatial arrangement of chromosomes. ..
  34. Bischoff M, Cseresnyes Z. Cell rearrangements, cell divisions and cell death in a migrating epithelial sheet in the abdomen of Drosophila. Development. 2009;136:2403-11 pubmed publisher
    ..Furthermore, our results suggest that an active force created by the histoblasts contributes to the formation of the adult epidermis. Finally, we show that histoblasts occasionally undergo apoptosis. ..
  35. Raisner R, Madhani H. Patterning chromatin: form and function for H2A.Z variant nucleosomes. Curr Opin Genet Dev. 2006;16:119-24 pubmed
    Although many histone variants are specific to higher eukaryotes, the H2A variant H2A.Z has been conserved during eukaryotic evolution. Genetic studies have demonstrated roles for H2A...
  36. Mehrotra S, McKim K. Temporal analysis of meiotic DNA double-strand break formation and repair in Drosophila females. PLoS Genet. 2006;2:e200 pubmed
    Using an antibody against the phosphorylated form of His2Av (gamma-His2Av), we have described the time course for the series of events leading from the formation of a double-strand break (DSB) to a crossover in Drosophila female meiotic ..
  37. Mavrich T, Jiang C, Ioshikhes I, Li X, Venters B, Zanton S, et al. Nucleosome organization in the Drosophila genome. Nature. 2008;453:358-62 pubmed publisher
    ..Here we produce a high-resolution reference map of H2A.Z and bulk nucleosome locations across the genome of the fly Drosophila melanogaster and compare it to that from ..
  38. van Daal A, White E, Gorovsky M, Elgin S. Drosophila has a single copy of the gene encoding a highly conserved histone H2A variant of the H2A.F/Z type. Nucleic Acids Res. 1988;16:7487-97 pubmed
    The Tetrahymena histone H2A variant designated hv1 is localized exclusively in the transcriptionally active macronucleus and is absent from the quiescent micronucleus (1)...
  39. Ivanovska I, Khandan T, Ito T, Orr Weaver T. A histone code in meiosis: the histone kinase, NHK-1, is required for proper chromosomal architecture in Drosophila oocytes. Genes Dev. 2005;19:2571-82 pubmed
    ..We describe a mutation in a recently identified Histone H2A kinase, nhk-1, in Drosophila that leads to female sterility due to defects in the formation of the meiotic ..
  40. McCaffrey R, St Johnston D, González Reyes A. Drosophila mus301/spindle-C encodes a helicase with an essential role in double-strand DNA break repair and meiotic progression. Genetics. 2006;174:1273-85 pubmed
    ..However, neither mei-W68 nor mei-41 rescue the defects in oocyte specification of mus301 mutants, suggesting that this helicase has another function in oocyte selection that is independent from its role in meiotic recombination. ..
  41. Kusch T, Florens L, Macdonald W, Swanson S, Glaser R, Yates J, et al. Acetylation by Tip60 is required for selective histone variant exchange at DNA lesions. Science. 2004;306:2084-7 pubmed
    Phosphorylation of the human histone variant H2A.X and H2Av, its homolog in Drosophila melanogaster, occurs rapidly at sites of DNA double-strand breaks...
  42. Klovstad M, Abdu U, Schupbach T. Drosophila brca2 is required for mitotic and meiotic DNA repair and efficient activation of the meiotic recombination checkpoint. PLoS Genet. 2008;4:e31 pubmed publisher
    ..In addition, Brca2 co-immunoprecipitates with the checkpoint protein Rad9, suggesting a direct role for Brca2 in the transduction of the meiotic recombination checkpoint signal. ..
  43. Kotadia S, Kao L, Comerford S, Jones R, Hammer R, Megraw T. PP2A-dependent disruption of centrosome replication and cytoskeleton organization in Drosophila by SV40 small tumor antigen. Oncogene. 2008;27:6334-46 pubmed publisher
    ..These analyses demonstrate the validity and utility of Drosophila as a model for viral oncoprotein function in vivo. ..
  44. Rogakou E, Boon C, Redon C, Bonner W. Megabase chromatin domains involved in DNA double-strand breaks in vivo. J Cell Biol. 1999;146:905-16 pubmed
    ..breaks introduced into mammalian cells by ionizing radiation results in the specific phosphorylation of histone H2AX on serine residue 139, yielding a specific modified form named gamma-H2AX...
  45. Leach T, Mazzeo M, Chotkowski H, Madigan J, Wotring M, Glaser R. Histone H2A.Z is widely but nonrandomly distributed in chromosomes of Drosophila melanogaster. J Biol Chem. 2000;275:23267-72 pubmed
    ..H2A.F/Z is a highly conserved family of H2A variants. H2Av, the H2A...
  46. van Daal A, Elgin S. A histone variant, H2AvD, is essential in Drosophila melanogaster. Mol Biol Cell. 1992;3:593-602 pubmed
    H2AvD, a Drosophila melanogaster histone variant of the H2A.Z class, is encoded by a single copy gene in the 97CD region of the polytene chromosomes...
  47. Clarkson M, Saint R. A His2AvDGFP fusion gene complements a lethal His2AvD mutant allele and provides an in vivo marker for Drosophila chromosome behavior. DNA Cell Biol. 1999;18:457-62 pubmed
    ..We show here that the fusion protein consists of functional GFP and functional histone His2AvD. The His2AvD portion of the fusion gene was shown to be functional by rescue of His2AvD mutant lethality...
  48. Pandey R, Heidmann S, Lehner C. Epithelial re-organization and dynamics of progression through mitosis in Drosophila separase complex mutants. J Cell Sci. 2005;118:733-42 pubmed
  49. Yu K, Saint R, Sullivan W. The Grapes checkpoint coordinates nuclear envelope breakdown and chromosome condensation. Nat Cell Biol. 2000;2:609-15 pubmed
    ..These results are likely to apply to vertebrate cells and suggest new strategies for destroying checkpoint-compromised cancer cells. ..
  50. Rong Y. Loss of the histone variant H2A.Z restores capping to checkpoint-defective telomeres in Drosophila. Genetics. 2008;180:1869-75 pubmed publisher
    The conserved histone variant H2A.Z fulfills many functions by being an integral part of the nucleosomes placed at specific regions of the genome...
  51. McCleland M, O Farrell P. RNAi of mitotic cyclins in Drosophila uncouples the nuclear and centrosome cycle. Curr Biol. 2008;18:245-54 pubmed publisher
    ..We suggest that high mitotic cyclin normally ensures that the centrosome cycle remains entrained to the nuclear cycle. ..
  52. Wells B, Johnston L. Maintenance of imaginal disc plasticity and regenerative potential in Drosophila by p53. Dev Biol. 2012;361:263-76 pubmed publisher
    ..find that after IR, p53 is required for imaginal disc cells to repair DNA, and in its absence the damage marker, ?-H2AX is persistently expressed...
  53. Stumpff J, Duncan T, Homola E, Campbell S, Su T. Drosophila Wee1 kinase regulates Cdk1 and mitotic entry during embryogenesis. Curr Biol. 2004;14:2143-8 pubmed
    ..These findings modify previous models about cell cycle regulation in syncytial embryos and demonstrate that Wee1 kinases can regulate mitotic entry in vivo during metazoan development even in cycles that lack a G2 phase. ..
  54. Hong A, Narbonne Reveau K, Riesgo Escovar J, Fu H, Aladjem M, Lilly M. The cyclin-dependent kinase inhibitor Dacapo promotes replication licensing during Drosophila endocycles. EMBO J. 2007;26:2071-82 pubmed
    ..Finally, genetic interaction studies suggest that dap functions to promote replication licensing in a subset of Drosophila mitotic cycles. ..