Affinage

H2AZ1

Histone H2A.Z · UniProt P0C0S5

Length
128 aa
Mass
13.6 kDa
Annotated
2026-06-10
100 papers in source corpus 50 papers cited in narrative 50 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

H2A.Z (H2AZ1/H2AFZ) is a conserved histone H2A variant deposited into nucleosomes flanking promoters and regulatory elements, where it tunes the chromatin template for transcription, replication, and genome stability (PMID:14645854, PMID:11081628, PMID:17392789). Its incorporation is catalyzed by the ATP-dependent SWR1/SRCAP-family remodeler, in which the Swr1/Domino ATPase drives histone exchange and the Swc2/YL1 (VPS72) subunit provides H2A.Z-specific recognition through a whip-like domain that engages the variant's hyperacidic patch and extended αC helix (PMID:14645854, PMID:16299513, PMID:26974124, PMID:26974126); single-molecule analysis resolves deposition into discrete priming, H2A loss, and dimer-release steps assisted by chaperones such as Chz1 and Nap1 (PMID:35263135, PMID:36396651, PMID:31107867, PMID:28883625). The reverse reaction is performed by INO80, which translocates uniquely at the H2A-H2B interface to evict H2A.Z-H2B dimers, working with the chaperone ANP32E (PMID:28604691, PMID:26142279). Structurally, H2A.Z's shorter C-terminus increases mobility of nucleosomal DNA termini and unwrapping while enabling more regular, condensed chromatin fibers, providing a basis for its context-dependent activating and repressive roles (PMID:34643712, PMID:32392318, PMID:24606920). At the +1 nucleosome H2A.Z lowers the barrier to elongating RNA Pol II (PMID:24606920), and it cooperates with Polycomb—being monoubiquitylated by RING1b to maintain repression of bivalent genes while antagonizing the bromodomain reader BRD2 (PMID:17636032, PMID:26804911)—and is mutually antagonistic with DNA methylation genome-wide (PMID:18815594). H2A.Z function is further tuned by post-translational modifications: acetylation by TIP60/KAT5 (notably at K7) and by KAT2A/GCN5 on H2A.Z.1 (recruiting BRD2 to promote RNAPII), monomethylation by SETD6 at K7, and dimethylation by SMYD3 at K101 that stabilizes the variant against ANP32E removal (PMID:35853868, PMID:31527837, PMID:23324626, PMID:27569210). Beyond transcription, H2A.Z scaffolds SUV420H1-mediated H4K20me2 to license early replication origins (PMID:31875854, PMID:37536340), supports chromosome segregation and centromere integrity, ES cell differentiation and lineage commitment, memory consolidation, and is required for proper nuclear reassembly after mitosis (PMID:34423893, PMID:18992931, PMID:25219850, PMID:32708675). Somatic and germline mutations in SRCAP-complex subunits cause defective H2A.Z deposition that drives epigenetic instability in uterine leiomyoma (PMID:34349258).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 2000 High

    Established that H2A.Z functions in transcriptional regulation through a chromatin pathway distinct from canonical remodeling/coactivator complexes, raising the question of how it is targeted to genes.

    Evidence Genetic epistasis and locus-specific ChIP at PHO5/GAL1 in S. cerevisiae

    PMID:11081628

    Open questions at the time
    • Did not identify the deposition machinery
    • Mechanism of locus-specific targeting unresolved
  2. 2003 High

    Identified the enzymatic basis of H2A.Z deposition, answering how the variant is placed into nucleosomes.

    Evidence Biochemical reconstitution of ATP-dependent H2A-for-H2A.Z exchange by the SWR1 complex plus in vivo ChIP in yeast

    PMID:14645854

    Open questions at the time
    • Which subunit confers H2A.Z specificity not yet defined
    • Step-by-step exchange mechanism unknown
  3. 2005 High

    Dissected SWR1 subunit roles and pinpointed H2A.Z C-terminal recognition, defining how the complex distinguishes the variant.

    Evidence Subunit pulldowns, in vitro exchange assays, and H2A.Z C-terminal mutagenesis

    PMID:16299513

    Open questions at the time
    • Atomic basis of Swc2-H2A.Z recognition not resolved
    • Chaperone contributions unclear
  4. 2007 High

    Mapped the genomic architecture of H2A.Z nucleosomes, establishing its hallmark positioning flanking nucleosome-free regions at promoters.

    Evidence Deep sequencing of ~322,000 H2A.Z nucleosomes genome-wide in yeast

    PMID:17392789

    Open questions at the time
    • Functional consequence of positioning at inactive genes unclear
    • No causal link to specific transcriptional outcomes
  5. 2007 Medium

    Revealed that H2A.Z carries distinguishing PTMs, with RING1b-mediated monoubiquitylation marking facultative heterochromatin.

    Evidence Fractionation, co-IP, immunofluorescence on the inactive X in mammalian cells

    PMID:17636032

    Open questions at the time
    • Functional consequence of ubiquitylation not yet tested
    • Site of modification not defined here
  6. 2008 High

    Connected H2A.Z to developmental gene regulation and Polycomb, and to mutual antagonism with DNA methylation, defining its epigenetic context.

    Evidence ChIP-seq/RNAi in mouse ES cells and reciprocal met1/pie1 mutant genomics in Arabidopsis

    PMID:18815594 PMID:18992931

    Open questions at the time
    • Molecular basis of H2A.Z-PcG interdependence unresolved
    • How methylation and H2A.Z exclude each other mechanistically unclear
  7. 2009 High

    Extended H2A.Z function to inducible mammalian enhancers/promoters, centromere silencing, antisense suppression, and identified acetylation sites that modulate nucleosome stability.

    Evidence ChIP-seq/RNAi at ERα targets (human), genetic epistasis at centromeres and antisense loci (S. pombe), and acetyl-site mass spec/NCP reconstitution

    PMID:19385636 PMID:19515975 PMID:19693008 PMID:19910462

    Open questions at the time
    • How acetylation state is read out in vivo unclear
    • Centromere role mechanism beyond CENP-C expression not defined
  8. 2013 High

    Identified specific writers placing methyl and acetyl marks on H2A.Z, linking modifications to differentiation and active transcription.

    Evidence In vitro enzymatic assays, mass spec, and ChIP for SETD6 (K7me1) and KAT2A/GCN5 (H2A.Z.1 acetylation recruiting BRD2)

    PMID:23324626 PMID:31527837

    Open questions at the time
    • Reader proteins for K7me1 not identified
    • Interplay among competing PTMs at a single residue unclear
  9. 2014 High

    Provided direct mechanistic evidence that H2A.Z lowers the Pol II elongation barrier and demonstrated an in vivo behavioral role in memory.

    Evidence Genome-wide NET-seq with H2A.Z depletion (yeast) and fear-conditioning ChIP/expression (mouse)

    PMID:24606920 PMID:25219850

    Open questions at the time
    • How H2A.Z structurally lowers the barrier not resolved here
    • Memory effect mechanism at gene-level undefined
  10. 2015 High

    Placed INO80/ANP32E as the H2A.Z removal pathway promoting homologous recombination, and identified BRD2 as an isoform-linked reader.

    Evidence Epistasis co-depletion with HR reporter assays and co-IP/ChIP-seq in human cells

    PMID:26051178 PMID:26142279

    Open questions at the time
    • Mechanistic basis of INO80 dimer eviction not yet structural
    • Isoform-specific BRD2 effects (H2A.Z.2 vs Z.1) need reconciling
  11. 2016 High

    Solved the structural basis of YL1/Swc2-mediated H2A.Z recognition and defined how methylation (SMYD3 K101me2) and ubiquitylation balance deposition versus removal and Polycomb repression.

    Evidence Crystal structures of YL1-H2A.Z-H2B (Drosophila and human), plus SMYD3 enzymatic/co-IP and H2A.Z.1 K3R3 mutagenesis with BRD2 rescue

    PMID:26804911 PMID:26974124 PMID:26974126 PMID:27569210

    Open questions at the time
    • How modifications are dynamically coordinated in cells unclear
    • Structural state during full SWR1 catalytic cycle still incomplete
  12. 2017 High

    Defined the INO80 eviction mechanism, identified the H2A.Z reader PWWP2A, and extended function to cotranscriptional splicing and redundant deposition chaperones.

    Evidence Single-molecule/biochemical INO80 translocation assays, H2A.Z interactome with domain mapping, S. pombe splicing genetics, and Nap1/Chz1 double-deletion analysis

    PMID:28446597 PMID:28604691 PMID:28645917 PMID:28883625

    Open questions at the time
    • How PWWP2A network directs downstream functions unclear
    • Splicing role mechanism beyond Prp16 pathway undefined
  13. 2018 Medium

    Expanded H2A.Z roles to Notch enhancer control, meiotic DSB formation, and gliogenesis via H3K56ac, showing functions across signaling, meiosis, and differentiation.

    Evidence ChIP/co-IP/knockdown at RBP-J enhancers, meiotic DSB and nuclear architecture assays (S. pombe), and ASF1a co-IP/ChIP in neural precursors

    PMID:29145618 PMID:29982651 PMID:29986055

    Open questions at the time
    • Direct vs indirect contributions to each pathway unclear
    • Single-lab findings without orthogonal confirmation
  14. 2019 High

    Established H2A.Z as a scaffold for replication-origin licensing and dissected chaperone bioavailability and tissue-specific deposition control.

    Evidence In vitro binding/ChIP-seq/nascent-strand sequencing for SUV420H1-H4K20me2-ORC1, Chz1 crystal structure, and Znhit1-YL1 phosphorylation in intestinal stem cells

    PMID:30842416 PMID:31107867 PMID:31875854

    Open questions at the time
    • How H2A.Z is restricted to early origins unclear
    • Coupling of deposition to replication timing not fully resolved
  15. 2020 High

    Linked H2A.Z structure to function—nucleosome unwrapping, CTCF regulation, isoform-specific compaction, and post-mitotic nuclear reassembly—while showing it is dispensable for transcription in non-dividing cells.

    Evidence MNase-X-ChIP-seq/CTCF CUT&RUN, cryo-EM of H2A.Z.2.2 nucleosome, cell-free nuclear reassembly, and in vivo muscle conditional knockout

    PMID:32109204 PMID:32266374 PMID:32392318 PMID:32708675 PMID:33073403

    Open questions at the time
    • Why H2A.Z is dispensable post-mitotically mechanistically unclear
    • Relationship between unwrapping and CTCF eviction incompletely defined
  16. 2021 High

    Provided structural and disease-level synthesis: cryo-EM of H2A.Z nucleosomes/fibers explaining dual roles, isoform division of labor in cell division, and SRCAP-mutation-driven H2A.Z loss in uterine leiomyoma.

    Evidence Cryo-EM with C-terminal mutagenesis, isoform-specific siRNA/imaging, Drosophila ZGA knockdown with HiC, and multi-omic primary-tumor sequencing

    PMID:34349258 PMID:34423893 PMID:34643712 PMID:34853314

    Open questions at the time
    • How fiber condensation reconciles with promoter activation unclear
    • Causal chain from SRCAP mutation to tumorigenesis incomplete
  17. 2022 High

    Resolved the temporal mechanism of SWR1 deposition and defined TIP60-dependent K7 acetylation, nutrient-responsive deacetylation, and acetylation-dependent epigenetic competence.

    Evidence Three-color single-molecule FRET and ensemble assays (yeast SWR1C), TIP60 knockout/ChIP, TORC1-Rpd3L-Ino80 deacetylation genetics, and Tip60-dependent bivalent gene activation

    PMID:35263135 PMID:35853868 PMID:36396651 PMID:36417913 PMID:36888706

    Open questions at the time
    • How discrete ATP steps couple to chaperone handoff in cells unclear
    • Whether deacetylation directly gates deposition in mammals unconfirmed
  18. 2023 High

    Defined SUV420H1 structural selectivity for H2A.Z nucleosomes and identified SENP5-mediated deSUMOylation regulating HR repair and radioresistance.

    Evidence Cryo-EM of SUV420H1-H2A.Z-nucleosome with mutagenesis, and SUMO-proteomics/co-IP/HR assays with PDO/PDX models

    PMID:37536340 PMID:37684630

    Open questions at the time
    • SUMO site(s) on H2A.Z not fully mapped
    • Therapeutic implications of SENP5 axis untested clinically
  19. 2024 Medium

    Showed that H2A.Z.1 acetylation status bidirectionally controls memory and couples to neuronal alternative splicing, linking specific PTMs to cognitive output.

    Evidence AAV-delivered acetyl-mimic/defective H2A.Z.1 mutants with fear conditioning and RNA/splicing analysis in mouse hippocampus

    PMID:38366138

    Open questions at the time
    • Reader machinery translating acetylation into splicing changes unknown
    • Single-lab in vivo finding without independent replication

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the full repertoire of H2A.Z post-translational modifications, isoform identity, and reader proteins is integrated to specify activating versus repressive outcomes at individual loci remains unresolved.
  • No unified model linking PTM combinations to context-specific function
  • Isoform-specific reader/effector logic only partially mapped
  • In vivo dynamics of deposition-eviction cycling at single loci undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 4 GO:0140110 transcription regulator activity 3 GO:0003677 DNA binding 2
Localization
GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 3 GO:0005694 chromosome 3
Pathway
R-HSA-74160 Gene expression (Transcription) 4 R-HSA-1266738 Developmental Biology 3 R-HSA-4839726 Chromatin organization 3 R-HSA-1640170 Cell Cycle 2 R-HSA-69306 DNA Replication 2 R-HSA-73894 DNA Repair 2 R-HSA-1643685 Disease 1
Complex memberships
INO80 complexSWR1/SRCAP complexTIP60/p400 complexnucleosome

Evidence

Reading pass · 50 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2003 The SWR1 complex, with Swr1 (a Swi2/Snf2-related ATPase) as its catalytic core, catalyzes ATP-dependent exchange of nucleosomal histone H2A for H2A.Z in nucleosome arrays. Swr1 is required for deposition of H2A.Z at specific chromosome locations in vivo. Biochemical reconstitution of histone exchange activity; in vivo ChIP; genetic analysis in S. cerevisiae Science High 14645854
2005 Within the SWR1 complex, Swc2 binds directly to H2A.Z and is essential for its transfer into nucleosomes. Swc6 and Arp6 are required for Swc2 association and nucleosome binding; Swc5 and Yaf9 are required for H2A.Z transfer but not for H2A.Z or nucleosome binding. The C-terminal alpha-helix of H2A.Z is crucial for recognition by SWR1. Biochemical dissection of SWR1 subunit interactions; pulldown/co-IP; in vitro histone exchange assay; mutagenesis of H2A.Z C-terminal helix Nature structural & molecular biology High 16299513
2007 H2A.Z nucleosomes occupy a single predominant rotational setting and multiple translational settings in the S. cerevisiae genome, flanking nucleosome-free regions at promoters of both active and inactive genes, with transcription factor binding sites rotationally exposed near nucleosome borders. Deep sequencing of 322,000 individual H2A.Z-containing nucleosomes (genome-wide nucleosome positioning) Nature High 17392789
2000 H2A.Z regulates gene transcription in S. cerevisiae; deletion of H2A.Z strongly increases the requirement for SNF/SWI and SAGA remodeling complexes (synthetic genetic interaction). H2A.Z is preferentially crosslinked to intergenic DNA at PHO5 and GAL1 loci, and this association changes with transcriptional activation, placing H2A.Z in a distinct pathway from SNF/SWI and SAGA for chromatin-based transcriptional regulation. Genetic epistasis (synthetic lethality/sick interactions); chromatin immunoprecipitation (ChIP) at specific loci in S. cerevisiae Cell High 11081628
2008 H2A.Z occupies promoters of developmentally important genes in mouse ES cells in a manner similar to Polycomb group protein Suz12. H2A.Z and PcG protein occupancy is interdependent at promoters (RNAi depletion of H2A.Z reduces PcG occupancy and vice versa), and H2A.Z is necessary for ES cell differentiation and lineage commitment. Genome-wide ChIP-seq; RNAi knockdown with gene expression and differentiation assays in mouse ES cells Cell High 18992931
2007 A fraction of mammalian H2A.Z is monoubiquitylated; on the inactive X chromosome the majority of H2A.Z is ubiquitylated. Monoubiquitylation of H2A.Z is mediated by the RING1b E3 ligase of the human Polycomb complex, distinguishing H2A.Z associated with facultative heterochromatin from euchromatin-associated H2A.Z. Biochemical fractionation; co-immunoprecipitation; immunofluorescence; identification of RING1b as the E3 ligase Molecular and cellular biology Medium 17636032
2008 DNA methylation and H2A.Z deposition are mutually antagonistic in Arabidopsis: regions of DNA methylation are quantitatively deficient in H2A.Z. Mutation of MET1 DNA methyltransferase causes losses of H2A.Z at sites gaining methylation and gains of H2A.Z at sites losing methylation; mutation of PIE1 (SWR1 complex subunit) leads to genome-wide hypermethylation. This demonstrates that DNA methylation excludes H2A.Z and H2A.Z protects genes from DNA methylation. ChIP-chip; genetic analysis of met1 and pie1 mutants with genome-wide methylation and H2A.Z profiling in Arabidopsis Nature High 18815594
2009 H2A.Z incorporation into promoter chromatin of estrogen receptor (ERα) target genes occurs upon gene induction in a cyclic pattern, mediated by the p400 complex. Cellular depletion of H2A.Z or p400 causes severe defects in estrogen signaling and estrogen-specific cell proliferation. H2A.Z incorporation allows nucleosomes to adopt preferential translational positions at the TFF1 promoter and is essential for estrogen-responsive enhancer function. ChIP-seq; RNAi knockdown; gene expression analysis; nucleosome positioning assay in human cells Genes & development High 19515975
2009 H2A.Z loaded at 5' ends of genes by the Swr1 complex containing a JmjC domain protein mediates suppression of antisense transcripts in fission yeast. H2A.Z is partially redundant with the Clr4-containing heterochromatin complex and RNAi component Ago1 in suppressing antisense transcripts that are normally degraded by the exosome. Genetic epistasis (H2A.Z/Clr4/Ago1 double and triple mutant analysis); ChIP; RNA analysis in S. pombe Nature High 19693008
2009 H2A.Z deposition in fission yeast requires a novel JmjC domain protein Msc1 as a component of the Swr1 complex. Loss of Msc1, Swr1, or H2A.Z results in loss of centromere silencing and defective chromosome segregation. H2A.Z is required for expression of the centromere protein CENP-C, and overexpression of CENP-C rescues centromere silencing defects associated with H2A.Z loss. Genetic analysis; ChIP; epistasis rescue experiments in S. pombe The Journal of biological chemistry Medium 19910462
2013 SETD6 lysine methyltransferase monomethylates H2A.Z at lysine 7 (H2AZK7me1). This modification increases upon differentiation of mouse embryonic stem cells; H2AZK7me1 and H3K27me3 co-occupy transcription start sites of differentiation marker genes. Depletion of Setd6 leads to cellular differentiation and compromised self-renewal in mESCs. In vitro methyltransferase assay; mass spectrometry; ChIP; siRNA knockdown in mESCs Epigenetics Medium 23324626
2015 H2A.Z.2 (encoded by H2AFV) interacts with BRD2 bromodomain protein. H2A.Z.2 controls transcriptional output of E2F target genes in melanoma; BRD2 binding to H2A.Z.2-regulated genes is dependent on H2A.Z.2 levels. H2A.Z.2 deficiency sensitizes melanoma cells to chemotherapy and targeted therapies. Co-immunoprecipitation; ChIP-seq; RNAi knockdown; integrated genomic analyses in melanoma cells Molecular cell Medium 26051178
2015 H2A.Z in human cells is rapidly removed from chromatin flanking DNA damage by the INO80 remodeling complex. The histone chaperone ANP32E also promotes homologous recombination and works in the same pathway as INO80 for H2A.Z removal. The HR defect in INO80- or ANP32E-depleted cells is rescued by co-depletion of H2A.Z, demonstrating that H2A.Z removal from chromatin is the primary function of INO80 and ANP32E in promoting HR. ChIP; epistasis rescue (co-depletion); siRNA knockdown; HR reporter assay in human cells EMBO reports High 26142279
2016 The crystal structure of Drosophila YL1 (Swc2) H2A.Z-binding domain (dYL1-Z) in complex with H2A.Z-H2B dimer at 1.9-Å resolution reveals a whip-like structure that wraps over H2A.Z-H2B. Preferential recognition is conferred by three residues in loop 2, the hyperacidic patch, and the extended αC helix of H2A.Z. This domain is essential for H2A.Z deposition in vivo and SRCAP(SWR1)-catalyzed replacement in vitro. Crystal structure (1.9 Å); in vitro histone replacement assay; in vivo deposition assay; mutagenesis Nature structural & molecular biology High 26974124
2016 YL1 is a specific H2A.Z-deposition chaperone in metazoans. Crystal structure at 2.7 Å of the human YL1-H2A.Z-H2B complex shows YL1 binding triggers extension of the H2A.Z αC helix and contacts the extended acidic patch and entire DNA-binding surface of H2A.Z-H2B. Only four amino acid substitutions in H2A are sufficient to create an H2A.Z-like interface recognized by YL1. Crystal structure (2.7 Å); biochemical binding assays; mutagenesis Nature structural & molecular biology High 26974126
2016 SMYD3 methyltransferase dimethylates H2A.Z.1 at lysine 101 (H2A.Z.1K101me2), which increases H2A.Z.1 stability by preventing binding to the removal chaperone ANP32E and facilitating interaction with histone H3. SMYD3 and H2A.Z.1K101me2 co-localize at the cyclin A1 promoter, activating its expression and G1-S progression. In vitro methyltransferase assay; co-immunoprecipitation; ChIP; cyclin A1 rescue in mouse model; microarray Cancer research Medium 27569210
2016 H2A.Z.1 monoubiquitylation (H2A.Z.1ub) at PRC1 target residues is required to maintain Polycomb binding and repress bivalent genes in mESCs. Loss of monoubiquitylation (K3R3 mutant) leads to de-repression of bivalent genes and faulty lineage commitment. Tandem bromodomain protein BRD2 is enriched in H2A.Z.1 chromatin and is gained at de-repressed promoters in H2A.Z.1(K3R3) mESCs; BRD2 inhibition restores gene silencing, revealing an antagonistic relationship between H2A.Z.1ub and BRD2. Mutagenesis (H2A.Z.1K3R3); ChIP; quantitative proteomics; BRD2 inhibition rescue experiment in mESCs Cell reports High 26804911
2017 Unlike other chromatin remodelers that translocate at the H3-H4 interface, INO80 translocates along DNA at the H2A-H2B interface of nucleosomes and persistently displaces DNA from H2A-H2B, generating DNA torsional strain near the entry site. This unique translocation mechanism drives both nucleosome mobilization and selective exchange of H2A.Z-H2B dimers for H2A-H2B without additional histone chaperones. INO80 translocates and mobilizes H2A.Z-containing nucleosomes more efficiently than H2A-containing nucleosomes. Biochemical translocation assays; single-molecule analysis; histone exchange assays in vitro; mutant analysis Nature communications High 28604691
2017 PWWP2A is a novel H2A.Z-nucleosome binding protein identified by H2A.Z interactome analysis. PWWP2A binds H2A.Z-containing nucleosomes via multivalent interaction (two internal regions for H2A.Z-specificity/nucleosome contact, PWWP domain for DNA binding). The C-terminal tail of H2A.Z mediates PWWP2A recruitment. PWWP2A depletion impairs cell proliferation via mitotic delay and causes neural crest cell defects in Xenopus. Co-immunoprecipitation interactome; domain mapping; genome-wide ChIP-seq; siRNA knockdown; Xenopus knockdown The EMBO journal Medium 28645917
2014 H2A.Z is actively exchanged in the hippocampus and cortex in response to fear conditioning. H2A.Z exchange mediates gene expression and restrains formation of recent and remote memory. H2A.Z acts as a negative regulator of hippocampal consolidation and systems consolidation. Fear conditioning behavioral paradigm; ChIP; gene expression analysis in mouse hippocampus and cortex Nature Medium 25219850
2019 H2A.Z-containing nucleosomes are enriched with H4K20me2 and bound ORC in HeLa cells. H2A.Z-nucleosomes bind directly to SUV420H1, promoting H4K20me2 deposition, which is required for ORC1 binding at replication origins. Depletion of H2A.Z reduces H4K20me2, ORC1 binding, and nascent-strand signals genome-wide, impairing early replication origin licensing and activation. In vitro binding assays; ChIP-seq; nascent strand sequencing; H2A.Z depletion in HeLa cells; genome-wide co-localization Nature High 31875854
2023 Cryo-EM structure of SUV420H1 bound to H2A.Z-nucleosome reveals direct interaction with H4 N-terminal tail (H4 residues 1-24 forming a lasso-shaped structure projecting H4K20 into the catalytic center), DNA, and nucleosome acidic patch. The SUV420H1 KR loop (residues 214-223) lies close to H2A.Z-specific residues D97/S98 and is crucial for preferential recognition of H2A.Z-nucleosome over canonical H2A-nucleosome. Cryo-EM structure; in vitro methyltransferase assay; mutagenesis; in vivo analysis Molecular cell High 37536340
2014 The +1 nucleosome is a major barrier to elongating RNAPII for essentially all genes. The extent of this barrier anticorrelates with enrichment of H2A.Z: depletion of H2A.Z from a nucleosome position results in a higher barrier to RNAPII. This shows H2A.Z incorporation at the +1 nucleosome reduces the transcriptional barrier. Genome-wide native elongating transcript sequencing (NET-seq) at nucleotide resolution; H2A.Z depletion in yeast Molecular cell High 24606920
2012 H2A.Z.2.2, an alternatively spliced variant of H2A.Z, binds to TIP60 and SRCAP chaperone complexes and is actively deposited into chromatin in a replication-independent manner. Its unique C-terminus causes major structural destabilization of nucleosomes as demonstrated by biochemical fractionation, FRAP, size exclusion chromatography, single-molecule FRET, and molecular dynamics simulations. Co-immunoprecipitation with TIP60/SRCAP complexes; FRAP; size exclusion chromatography; single-molecule FRET; molecular dynamics simulations; deletion mutant analysis Nucleic acids research High 22467210
2009 Acetylation sites of vertebrate H2A.Z (K4, K7, K11, K13, K15) were identified from chicken erythrocytes. H2A.Z stabilizes the nucleosome core particle (NCP) regardless of its acetylation state. Acetylated H2A.Z variants act synergistically with other acetylated core histones to alter NCP conformation. Heteromorphic NCPs containing both H2A.Z and H2A are slightly destabilized only in the presence of acetylation. Mass spectrometry (acetylation site identification); NCP reconstitution with acetylated histones and H2A.Z(K/Q) acetylation mimic mutants; biochemical stability assays Biochemistry Medium 19385636
2019 Znhit1, a subunit of the SRCAP complex, incorporates H2A.Z into TSS regions of genes governing Lgr5+ intestinal stem cell fate (Lgr5, Tgfb1, Tgfbr2). Mechanistically, Znhit1 promotes H2A.Z incorporation by controlling phosphorylation of the H2A.Z chaperone YL1, enhancing YL1-H2A.Z interaction. Conditional knockout mouse; ChIP-seq; co-immunoprecipitation; phosphorylation analysis; intestinal stem cell functional assays Nature communications Medium 30842416
2018 H2A.Z deposition and acetylation by the TRRAP/p400/Tip60 complex (which physically associates with transcription factor RBP-J at Notch-dependent enhancers) is required for tight control of canonical Notch target gene activation. Tip60 acetylates H2A.Z when targeted to RBP-J-bound enhancers, upregulating Notch target genes. Drosophila homologs of Tip60, p400, and H2A.Z modulate Notch signaling in vivo. ChIP; co-immunoprecipitation; siRNA knockdown; in vivo Drosophila genetic analysis; gene expression assays Nucleic acids research Medium 29986055
2013 KAT2A (GCN5) acetylates H2A.Z.1 at promoters of transactivated genes. The DNA repair complex XPC-RAD23-CEN2 interacts with both H2A.Z and KAT2A, recruiting KAT2A to promoters to license H2A.Z.1 acetylation. KAT2A selectively acetylates H2A.Z.1 but not H2A.Z.2 in vitro; alanine-14 in H2A.Z.2 inhibits KAT2A activity. Acetyl-H2A.Z.1 recruits BRD2 to promote RNAPII recruitment. In vitro acetyltransferase assay; co-immunoprecipitation; ChIP; mutagenesis (non-acetylable H2A.Z.1); RNAPII recruitment assay Nature chemical biology High 31527837
2022 TIP60 (KAT5) is essential for acetylation of H2A.Z specifically at lysine 7 in human and mouse cells. Loss of TIP60 causes complete cell growth arrest with failure of chromosome alignment during mitosis. This growth arrest is independent of p53, INK4A, and ARF tumor suppressors. Inducible Cre-recombinase and CRISPR/Cas9 deletion of TIP60; ChIP; RNA-seq; cell cycle analysis; live imaging of mitosis Cell death & disease Medium 35853868
2022 Tip60 acetyltransferase acetylates H2A.Z, and loss of H2A.Z acetylation interferes with H3K4me3 deposition and activation of bivalent, lineage-restricted genes during neuronal fate specification. H2A.Z acetylation does not affect chromatin accessibility but is required for epigenetic competence (bivalent chromatin activation) during cell lineage transition. Proteomics; genetic depletion of Tip60; ChIP-seq; ATAC-seq; RNA-seq; neuronal differentiation assays Molecular cell Medium 36417913
2020 H2A.Z nucleosomes are more enriched in unwrapped states compared with canonical nucleosomes in mouse ES cells, as measured by MNase-X-ChIP-seq. Depletion of H2A.Z results in decreased unwrapping of H3.3 nucleosomes and increased CTCF binding, demonstrating that H2A.Z regulates nucleosome unwrapping in vivo and that its transcriptional and CTCF-regulatory functions correlate with nucleosome unwrapping states. MNase-X-ChIP-seq; re-ChIP; CTCF CUT&RUN; H2A.Z depletion in mouse ES cells Nucleic acids research Medium 32392318
2021 Cryo-EM structures of H2A.Z-containing nucleosomes and chromatin fibers show that H2A.Z incorporation increases mobility of DNA termini in nucleosomes (due to its shorter C-terminus) while simultaneously enabling nucleosome arrays to form a more regular and condensed chromatin fiber. This structural basis provides a mechanistic explanation for H2A.Z's dual activating and repressive transcriptional roles. Cryo-EM structure of nucleosome and chromatin fiber; deletion mutant analysis of H2A.Z C-terminus Nucleic acids research High 34643712
2020 Cryo-EM structure of H2A.Z.2.2-containing nucleosome (Z.2.2-NCP) shows it compacts ~125 bp of DNA (less than canonical ~147 bp). Nineteen H2A.Z.2.2-specific residues including a ROF ('regulating-octamer-folding') sequence are responsible for nucleosome opening and also required for SWR1-dependent histone replacement. Cryo-EM structure; MNase digestion assay; mutagenesis of ROF sequence; SWR1-dependent replacement assay The EMBO journal High 33073403
2022 Single-molecule FRET analysis of SWR1-mediated H2A.Z deposition reveals three ATP-dependent phases: (1) a priming step involving transient DNA unwrapping and histone octamer deformations; (2) rapid loss of H2A-H2B; (3) release of displaced H2A-H2B from the SWR1-nucleosome complex, facilitated by histone chaperones. SWR1 intrinsically senses long-linker DNA to preferentially exchange H2A.Z on the distal face of asymmetrically positioned nucleosomes. Three-color single-molecule FRET; ensemble biochemical assays; real-time single nucleosome remodeling imaging Science advances High 35263135
2022 SWR1C (yeast SWR1 complex) uses multiple ATP-dependent steps for H2A.Z deposition, identified by single-molecule and ensemble methods: an initial priming step (transient DNA unwrapping and octamer deformations), rapid H2A loss, and subsequent H2A/H2B release from the H2A.Z nucleosomal product. Rates of priming and H2A/H2B dimer release are sensitive to ATP concentration. Single-molecule real-time analysis of nucleosome remodeling; ensemble histone exchange assays; ATP concentration titrations Nature communications High 36396651
2019 The crystal structure of the Chz1-H2A.Z-H2B ternary complex reveals that Chz1 uses two distinct structural domains: the middle region (Chz1-M) directly interacts with H2A.Z-specific residues Gly98 and Ala57 for modest preferential binding; the C-terminal region (Chz1-C) harbors a DEF/Y motif engaging an arginine finger and hydrophobic pocket in H2A.Z-H2B, enhancing binding preference. Chz1 facilitates SWR1-mediated H2A.Z deposition by controlling bioavailability of free H2A.Z. Crystal structure; biochemical binding assays; mutagenesis; in vitro SWR1 histone deposition assay PLoS biology High 31107867
2017 Nap1 and Chz1 have redundant roles as histone chaperones for H2A.Z deposition in S. cerevisiae. They use different H2A.Z surface residues for interaction. Loss of either Nap1 or Chz1 alone causes mild H2A.Z deposition defects, but deletion of both results in significant reduction of H2A.Z at promoters and heterochromatin spreading. Structural modeling; molecular dynamics simulations; mutagenesis; ChIP; genetic double deletion analysis in S. cerevisiae Scientific reports Medium 28883625
2021 H2A.Z.1 and H2A.Z.2 have non-redundant functions in cell division. H2A.Z.1 regulates expression of cell cycle genes (Myc, Ki-67) and its depletion leads to G1 arrest and cellular senescence. H2A.Z.2, in a transcription-independent manner, is essential for centromere integrity and sister chromatid cohesion regulation, playing a key role in chromosome segregation. Isoform-specific siRNA depletion; cell cycle analysis; live-cell imaging; ChIP-seq; RNA-seq in human cells EMBO reports Medium 34423893
2021 H2A.Z.1 and H2A.Z.2 have specific interactors (identified by mass spectrometry) that can mediate functional antagonism. They can replace each other at transcription start sites and regulate both distinct and overlapping gene sets in a context-dependent manner. The balance between the two isoforms at promoters critically regulates specific gene expression. Endogenously-tagged protein immunoprecipitation; mass spectrometry; RNA-seq; ChIP-seq in human cells eLife Medium 32109204
2023 SENP5 deSUMOylates H2A.Z, and this modification regulates homologous recombination-mediated DNA damage repair. The SUMOylation balance of H2A.Z modulates HR repair and cancer radioresistance. H2A.Z was identified as a SENP5 deSUMOylation substrate by SUMO-proteomic mass spectrometry and co-immunoprecipitation. SUMO-proteomic mass spectrometry; co-immunoprecipitation; HR reporter assay; siRNA knockdown; PDO and PDX models Journal of experimental & clinical cancer research Medium 37684630
2023 TORC1 activates the Rpd3L histone deacetylase complex to deacetylate H2A.Z, blocking its deposition into chromatin at autophagy-related gene promoters. Simultaneously, Rpd3L deacetylates the chromatin remodeler Ino80 at K929, stabilizing it and enabling H2A.Z eviction from autophagy genes. This TORC1-Rpd3L-H2A.Z axis represses autophagy-related gene transcription under nutrient-rich conditions. Biochemical deacetylase assays; co-immunoprecipitation; ChIP; genetic epistasis; rapamycin treatment in yeast Science advances Medium 36888706
2020 VPS72/YL1 mediates H2A.Z deposition required for nuclear reassembly after mitosis. Depletion of VPS72 or H2A.Z by siRNA or in vitro depletion results in malformed and nonfunctional nuclei. VPS72 functions outside of the SRCAP and EP400 remodeling complexes for this specific H2A.Z deposition activity. Cell-free nuclear reassembly assay; siRNA knockdown; live-cell imaging; biochemical fractionation Cells Medium 32708675
2017 H2A.Z promotes cotranscriptional splicing of suboptimal introns in S. pombe. H2A.Z and Swr1 ATPase are required during temperature stress for efficient splicing of introns with nonconsensus splice sites. Mutating affected splice sites to consensus suppresses the H2A.Z requirement for splicing of that intron. Overexpression of splicing ATPase Prp16 suppresses growth and splicing defects of H2A.Z mutants, placing H2A.Z upstream of Prp16 in a pathway promoting weak intron recognition. Epistatic miniarray profiles (EMAP); splicing microarray; splice site mutagenesis; genetic suppression by Prp16 overexpression in S. pombe Genes & development Medium 28446597
2021 H2A.Z depletion in terminally differentiated post-mitotic muscle cells in vivo shows that H2A.Z is dispensable for maintaining or activating transcription in the absence of cell division, as shown by ChIP-seq, RNA-seq, and ATAC-seq. H2A.Z enrichment at active promoters is a marker but not a driver of transcription in post-mitotic cells. In vivo conditional H2A.Z knockout in post-mitotic muscle; ChIP-seq; RNA-seq; ATAC-seq Nucleic acids research Medium 32266374
2021 In Drosophila embryos, H2A.Z enrichment at transcription start sites precedes ZGA and RNA Pol II loading. In vivo knockdown of Domino (H2A.Z chaperone/ATPase) reduces H2A.Z deposition at TSS, causes global downregulation of housekeeping genes at ZGA, and compromises establishment of 3D chromatin structure. GRO-seq; HiC; ChIP-seq; in vivo RNAi knockdown in Drosophila embryos Nature communications Medium 34853314
2018 H2A.Z promotes meiotic DNA double-strand break formation in fission yeast. In H2A.Z-lacking mutants, multiple proteins involved in DSB formation (including Spo11 homolog and its regulators) show reduced chromatin association, and nuclei are more compact. H2A.Z promotes DSB formation by modulating chromosome architecture to enhance interaction between DSB-related proteins and cohesin-loaded chromatin. ChIP; meiotic DSB assay; nuclear architecture analysis; genetic analysis in S. pombe Nucleic acids research Medium 29145618
2018 H2A.Z.1 regulates gliogenesis by cooperating with histone chaperone ASF1a to regulate H3K56 acetylation (H3K56ac), which in turn directly affects expression of FOLR1 (folate receptor 1). FOLR1 participates in gliogenesis through the JAK-STAT signaling pathway. siRNA knockdown; ChIP; RNA-seq; co-immunoprecipitation with ASF1a; neural precursor cell differentiation assays Nucleic acids research Medium 29982651
2015 The DREAM complex (containing C. elegans lin-35/Rb homolog) promotes high gene-body H2A.Z (HTZ-1) levels at target genes. In lin-35 mutants, DREAM targets show reduced gene body H2A.Z and increased expression. Many DREAM targets are also upregulated in htz-1/H2A.Z mutants, indicating that gene body H2A.Z plays a repressive role at DREAM target genes. ChIP-seq; genetic epistasis (lin-35 and htz-1 mutants); RNA-seq in C. elegans Genes & development Medium 25737279
2024 The role of H2A.Z.1 in memory depends on its acetylation status: acetyl-mimic H2A.Z.1 (K4/7/11Q) improves memory, while acetyl-defective H2A.Z.1 (K4/7/11A) impairs memory. H2A.Z.1KA preferentially affects genes involved in synaptic function. H2A.Z.1 depletion or expression of lysine mutants influences alternative splicing of neuronal genes, revealing that H2A.Z.1 regulates both gene expression and splicing in neurons through post-translational modifications. AAV-mediated overexpression of acetyl-mimic and acetyl-defective H2A.Z.1 mutants; fear conditioning behavioral assays; RNA-seq; splicing analysis in mouse hippocampus Neuropsychopharmacology Medium 38366138
2021 Somatic mutations in SRCAP complex subunit genes cause defective H2A.Z deposition in uterine leiomyomas. In these tumors, H2A.Z loss at TSS is associated with increased chromatin accessibility and upregulation of bivalent embryonic stem cell genes. Germline mutations in SRCAP members YEATS4 and ZNHIT1 predispose to uterine leiomyoma through epigenetic instability caused by deficient H2A.Z deposition. Whole-genome sequencing; ATAC-seq; ChIP-seq; HiChIP; RNA-seq of primary tumor tissues Nature High 34349258

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2003 ATP-driven exchange of histone H2AZ variant catalyzed by SWR1 chromatin remodeling complex. Science (New York, N.Y.) 1032 14645854
2002 Histone H2A variants H2AX and H2AZ. Current opinion in genetics & development 592 11893489
2007 Translational and rotational settings of H2A.Z nucleosomes across the Saccharomyces cerevisiae genome. Nature 548 17392789
2008 Histone H2A.Z and DNA methylation are mutually antagonistic chromatin marks. Nature 438 18815594
2014 Nucleosomes are context-specific, H2A.Z-modulated barriers to RNA polymerase. Molecular cell 302 24606920
2008 H2AZ is enriched at polycomb complex target genes in ES cells and is necessary for lineage commitment. Cell 281 18992931
2000 Histone H2A.Z regulats transcription and is partially redundant with nucleosome remodeling complexes. Cell 269 11081628
2019 The histone variant H2A.Z in gene regulation. Epigenetics & chromatin 252 31200754
2008 H2A.Z: view from the top. Structure (London, England : 1993) 215 18275809
2005 Swc2 is a widely conserved H2AZ-binding module essential for ATP-dependent histone exchange. Nature structural & molecular biology 205 16299513
2015 Histone Variant H2A.Z.2 Mediates Proliferation and Drug Sensitivity of Malignant Melanoma. Molecular cell 164 26051178
2007 Monoubiquitylation of H2A.Z distinguishes its association with euchromatin or facultative heterochromatin. Molecular and cellular biology 149 17636032
2014 Histone H2A.Z subunit exchange controls consolidation of recent and remote memory. Nature 147 25219850
2019 H2A.Z facilitates licensing and activation of early replication origins. Nature 146 31875854
2009 Histone H2A.Z is essential for estrogen receptor signaling. Genes & development 130 19515975
2015 Removal of H2A.Z by INO80 promotes homologous recombination. EMBO reports 124 26142279
2009 Histone H2A.Z cooperates with RNAi and heterochromatin factors to suppress antisense RNAs. Nature 121 19693008
2017 INO80 exchanges H2A.Z for H2A by translocating on DNA proximal to histone dimers. Nature communications 120 28604691
2009 The beauty of being a variant: H2A.Z and the SWR1 complex in plants. Molecular plant 112 19825639
2010 Changes in H2A.Z occupancy and DNA methylation during B-cell lymphomagenesis. Genome research 104 20709945
2009 Characterization of the histone H2A.Z-1 and H2A.Z-2 isoforms in vertebrates. BMC biology 91 20003410
2012 H2A.Z.2.2 is an alternatively spliced histone H2A.Z variant that causes severe nucleosome destabilization. Nucleic acids research 90 22467210
2010 Reconciling the positive and negative roles of histone H2A.Z in gene transcription. Epigenetics 90 20364108
2013 Precise deposition of histone H2A.Z in chromatin for genome expression and maintenance. Biochimica et biophysica acta 88 24459731
2021 Deficient H2A.Z deposition is associated with genesis of uterine leiomyoma. Nature 86 34349258
2006 Patterning chromatin: form and function for H2A.Z variant nucleosomes. Current opinion in genetics & development 83 16503125
2016 Structural basis of H2A.Z recognition by SRCAP chromatin-remodeling subunit YL1. Nature structural & molecular biology 82 26974124
2009 Acetylation of vertebrate H2A.Z and its effect on the structure of the nucleosome. Biochemistry 81 19385636
2015 H2A.Z: a molecular rheostat for transcriptional control. F1000prime reports 79 25705384
2018 Learning and Age-Related Changes in Genome-wide H2A.Z Binding in the Mouse Hippocampus. Cell reports 76 29386101
2013 Gene dysregulation by histone variant H2A.Z in bladder cancer. Epigenetics & chromatin 71 24279307
2016 H2A.Z controls the stability and mobility of nucleosomes to regulate expression of the LH genes. Nature communications 70 27653784
2016 Molecular basis and specificity of H2A.Z-H2B recognition and deposition by the histone chaperone YL1. Nature structural & molecular biology 68 26974126
2023 Histone variant H2A.Z modulates nucleosome dynamics to promote DNA accessibility. Nature communications 67 36765119
2005 Histone H2AZ dimerizes with a novel variant H2B and is enriched at repetitive DNA in Trypanosoma brucei. Journal of cell science 65 16303849
2021 The H2A.Z-nuclesome code in mammals: emerging functions. Trends in genetics : TIG 64 34702577
2016 Effects of MacroH2A and H2A.Z on Nucleosome Dynamics as Elucidated by Molecular Dynamics Simulations. Biophysical journal 59 26789756
2013 SETD6 monomethylates H2AZ on lysine 7 and is required for the maintenance of embryonic stem cell self-renewal. Epigenetics 57 23324626
2016 H2A.Z.1 Monoubiquitylation Antagonizes BRD2 to Maintain Poised Chromatin in ESCs. Cell reports 55 26804911
2021 Histone variant H2A.Z regulates zygotic genome activation. Nature communications 52 34853314
2015 The DREAM complex promotes gene body H2A.Z for target repression. Genes & development 52 25737279
2009 Histone variant H2A.Z regulates centromere silencing and chromosome segregation in fission yeast. The Journal of biological chemistry 50 19910462
2021 Structural basis of chromatin regulation by histone variant H2A.Z. Nucleic acids research 49 34643712
2016 SMYD3-Mediated H2A.Z.1 Methylation Promotes Cell Cycle and Cancer Proliferation. Cancer research 49 27569210
2019 Znhit1 controls intestinal stem cell maintenance by regulating H2A.Z incorporation. Nature communications 48 30842416
2009 Transcriptional and epigenetic functions of histone variant H2A.Z. Biochemistry and cell biology = Biochimie et biologie cellulaire 48 19234520
2017 Multivalent binding of PWWP2A to H2A.Z regulates mitosis and neural crest differentiation. The EMBO journal 45 28645917
2018 Histone variant H2A.Z deposition and acetylation directs the canonical Notch signaling response. Nucleic acids research 44 29986055
2020 Histone variant H2A.Z regulates nucleosome unwrapping and CTCF binding in mouse ES cells. Nucleic acids research 42 32392318
2020 Structural basis of nucleosome dynamics modulation by histone variants H2A.B and H2A.Z.2.2. The EMBO journal 42 33073403
2021 Multiple roles of H2A.Z in regulating promoter chromatin architecture in human cells. Nature communications 41 33953180
2012 H2A.Z-dependent crosstalk between enhancer and promoter regulates cyclin D1 expression. Oncogene 41 23108396
2010 Histone variant H2A.Z can serve as a new target for breast cancer therapy. Current medicinal chemistry 41 20666725
2021 Non-redundant functions of H2A.Z.1 and H2A.Z.2 in chromosome segregation and cell cycle progression. EMBO reports 40 34423893
2012 DNA methylation, H2A.Z, and the regulation of constitutive expression. Cold Spring Harbor symposia on quantitative biology 40 23250988
2020 Integrated analysis of H2A.Z isoforms function reveals a complex interplay in gene regulation. eLife 39 32109204
2020 Distinct roles for H4 and H2A.Z acetylation in RNA transcription in African trypanosomes. Nature communications 39 32198348
2020 AtINO80 represses photomorphogenesis by modulating nucleosome density and H2A.Z incorporation in light-related genes. Proceedings of the National Academy of Sciences of the United States of America 38 33318175
2022 Hierarchical Accumulation of Histone Variant H2A.Z Regulates Transcriptional States and Histone Modifications in Early Mammalian Embryos. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 37 35717671
2021 H2A.Z overexpression suppresses senescence and chemosensitivity in pancreatic ductal adenocarcinoma. Oncogene 37 33627784
2023 Coordinated histone variant H2A.Z eviction and H3.3 deposition control plant thermomorphogenesis. The New phytologist 35 36647799
2018 H2A.Z-containing nucleosomes are evicted to activate AtMYB44 transcription in response to salt stress. Biochemical and biophysical research communications 33 29649476
2013 TIP48/Reptin and H2A.Z requirement for initiating chromatin remodeling in estrogen-activated transcription. PLoS genetics 33 23637611
2022 Tip60-mediated H2A.Z acetylation promotes neuronal fate specification and bivalent gene activation. Molecular cell 31 36417913
2022 E2F and STAT3 provide transcriptional synergy for histone variant H2AZ activation to sustain glioblastoma chromatin accessibility and tumorigenicity. Cell death and differentiation 28 35058574
2022 Coordinated DNA and histone dynamics drive accurate histone H2A.Z exchange. Science advances 28 35263135
2014 Dynamics of histone variant H3.3 and its coregulation with H2A.Z at enhancers and promoters. Nucleus (Austin, Tex.) 28 24637397
2023 SENP5 promotes homologous recombination-mediated DNA damage repair in colorectal cancer cells through H2AZ deSUMOylation. Journal of experimental & clinical cancer research : CR 27 37684630
2020 Variation on a theme: Evolutionary strategies for H2A.Z exchange by SWR1-type remodelers. Current opinion in cell biology 27 33217681
2019 The H2A.Z histone variant integrates Wnt signaling in intestinal epithelial homeostasis. Nature communications 27 31015444
2014 Histone H2A.Z deregulation in prostate cancer. Cause or effect? Cancer metastasis reviews 27 24398858
2020 H2A.Z is dispensable for both basal and activated transcription in post-mitotic mouse muscles. Nucleic acids research 26 32266374
2021 INO80 promotes H2A.Z occupancy to regulate cell fate transition in pluripotent stem cells. Nucleic acids research 25 34139016
2019 DNA repair complex licenses acetylation of H2A.Z.1 by KAT2A during transcription. Nature chemical biology 25 31527837
2018 H2A.Z regulates tumorigenesis, metastasis and sensitivity to cisplatin in intrahepatic cholangiocarcinoma. International journal of oncology 25 29532867
2022 H2A.Z's 'social' network: functional partners of an enigmatic histone variant. Trends in biochemical sciences 24 35606214
2022 Loss of TIP60 (KAT5) abolishes H2AZ lysine 7 acetylation and causes p53, INK4A, and ARF-independent cell cycle arrest. Cell death & disease 24 35853868
2018 The histone variant H2A.Z promotes initiation of meiotic recombination in fission yeast. Nucleic acids research 24 29145618
2019 Structural insights into histone chaperone Chz1-mediated H2A.Z recognition and histone replacement. PLoS biology 22 31107867
2011 Molecular cloaking of H2A.Z on mortal DNA chromosomes during nonrandom segregation. Stem cells (Dayton, Ohio) 22 21905168
2020 VPS72/YL1-Mediated H2A.Z Deposition Is Required for Nuclear Reassembly after Mitosis. Cells 21 32708675
2017 The histone variant H2A.Z promotes splicing of weak introns. Genes & development 21 28446597
2010 H2A.Z (Htz1) controls the cell-cycle-dependent establishment of transcriptional silencing at Saccharomyces cerevisiae telomeres. Genetics 21 20980239
2023 The TORC1 activates Rpd3L complex to deacetylate Ino80 and H2A.Z and repress autophagy. Science advances 20 36888706
2017 Recognition of hyperacetylated N-terminus of H2AZ by TbBDF2 from Trypanosoma brucei. The Biochemical journal 20 29025975
2017 H2A O-GlcNAcylation at serine 40 functions genomic protection in association with acetylated H2AZ or γH2AX. Epigenetics & chromatin 20 29084613
2020 Vital and Distinct Roles of H2A.Z Isoforms in Hepatocellular Carcinoma. OncoTargets and therapy 18 32547065
2018 H2A.Z.1 crosstalk with H3K56-acetylation controls gliogenesis through the transcription of folate receptor. Nucleic acids research 18 29982651
2023 Structural insight into H4K20 methylation on H2A.Z-nucleosome by SUV420H1. Molecular cell 17 37536340
2022 GAS41 mediates proliferation and GEM chemoresistance via H2A.Z.2 and Notch1 in pancreatic cancer. Cellular oncology (Dordrecht, Netherlands) 17 35503594
2023 The H2A.Z and NuRD associated protein HMG20A controls early head and heart developmental transcription programs. Nature communications 16 36709316
2022 H2A.Z deposition by SWR1C involves multiple ATP-dependent steps. Nature communications 16 36396651
2017 Genome-wide identification of histone H2A and histone variant H2A.Z-interacting proteins by bPPI-seq. Cell research 16 28862252
2017 Redundant Functions for Nap1 and Chz1 in H2A.Z Deposition. Scientific reports 16 28883625
2024 Role of the histone variant H2A.Z.1 in memory, transcription, and alternative splicing is mediated by lysine modification. Neuropsychopharmacology : official publication of the American College of Neuropsychopharmacology 15 38366138
2022 Contribution of the histone variant H2A.Z to expression of responsive genes in plants. Seminars in cell & developmental biology 15 35474148
2020 Histone H2A.Z is required for androgen receptor-mediated effects on fear memory. Neurobiology of learning and memory 15 32916283
2018 Metformin alters H2A.Z dynamics and regulates androgen dependent prostate cancer progression. Oncotarget 15 30651935
2015 H2A.Z marks antisense promoters and has positive effects on antisense transcript levels in budding yeast. BMC genomics 15 25765960
2014 Yeast X-chromosome-associated protein 5 (Xap5) functions with H2A.Z to suppress aberrant transcripts. EMBO reports 15 24957674

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