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HistonesHistone DeacetylasesHistone Deacetylase InhibitorsHistone AcetyltransferasesAcetylationAmino Acid SequenceMolecular Sequence DataChromatinHistone Deacetylase 1MethylationHistone CodeHistone Deacetylase 2Histone DemethylasesHistone ChaperonesNucleosomesBinding SitesModels, MolecularProtein BindingHydroxamic AcidsHistone-Lysine N-MethyltransferaseProtein Structure, TertiaryMutagenesis, Site-DirectedBase SequenceProtein ConformationMutationSequence Homology, Amino AcidEpigenesis, GeneticTranscription, GeneticKineticsChromatin Assembly and DisassemblyJumonji Domain-Containing Histone DemethylasesProtein Processing, Post-TranslationalProtein Structure, SecondaryAcetyltransferasesSubstrate SpecificityStructure-Activity RelationshipSequence AlignmentCell LineAmino Acid SubstitutionDNAProtein MethyltransferasesRecombinant ProteinsSaccharomyces cerevisiae ProteinsChromatin ImmunoprecipitationAmino AcidsTranscription FactorsRepressor ProteinsNuclear ProteinsCysteinePeptide FragmentsDNA-Binding ProteinsSaccharomyces cerevisiaePhosphorylationCloning, MolecularEscherichia colip300-CBP Transcription FactorsCrystallography, X-RayHeLa CellsChromosomal Proteins, Non-HistoneGene SilencingPeptidesConserved SequenceDrug ResiduesHeterochromatinElectrophoresis, Polyacrylamide GelRecombinant Fusion ProteinsCatalytic DomainGene Expression RegulationDNA MethylationCatalysisHistidineAmino Acid MotifsPesticide ResiduesSerineLysineCircular DichroismMagnetic Resonance SpectroscopyCattleButyratesArginineEnzyme InhibitorsChickensRNA, MessengerMutagenesisMethyltransferasesProtamine KinaseDNA PrimersProtein FoldingProteinsPlasmidsAlanineTransfectionSin3 Histone Deacetylase and Corepressor ComplexMicrococcal NucleaseTrypsinBacterial ProteinsDimerizationCell Line, TumorHydrogen-Ion ConcentrationPoint Mutation
MethylationAcetylationProteinsLysine residues of histonesChromatin structurePeptidesTranscriptionalPhosphorylationEnzymesPeptideTranscriptionModificationsSpecific histoneGenesTailsDeacetylase InhibitorsGene expressionHDAcIsoformsAcetyltransferaseInhibitorsDeacetylasesRepressiveMethyltransferasesBindsInhibitionH3K4ModificationNucleosomeMammalianReplication independentBiochemicalMRNAOrthologsPolyclonalAmino acidsSpecificityUbiquitinationVariantMutationsChromosomeStructuralVariantsDistinctRecognizeAlterationsInteractionsRapamycinDrosophilaAsymmetricMolecularActivatesLocalization
Methylation19
  • Here, we summarize the different types of epigenetic enzymes which target corresponding protein domains, emphasize DNA methylation , histone modifications, and microRNA-mediated cooperation with epigenetic modification, and highlight recent achievements in developing targets for epigenetic inhibitor therapy. (frontiersin.org)
  • Protein methylation at lysine residues modulates chromatin structure, affects gene expression and mammalian development. (europa.eu)
  • Methylation of histones does not usually change as much as phosphorylation does. (news-medical.net)
  • Some methylation alterations of H3 histones occur in the M phase, whereas many other H3s do not change noticeably. (news-medical.net)
  • Histone methylation plays an important role in the regulation of chromatin structure and function. (eurogentec.com)
  • Dot1L mediated histone H3 lysine79 methylation is essential to meiosis progression in mouse oocytes. (nel.edu)
  • Wang X, Gao W, Ma X, Wang X, Song C, Huang X, Liu H. Dot1L mediated histone H3 lysine79 methylation is essential to meiosis progression in mouse oocytes. (nel.edu)
  • Histone methylation by the Drosophila epigenetic transcriptional regulator Ash1. (wikigenes.org)
  • Methylation of this fourth amino acid residue from the N-terminus of histone H3 is one of the most studied histone modifications, and with good reason: it's tightly associated with the promoters of active genes. (epigenie.com)
  • While this is still an active area of research, it is clear that specific histone methylation states regulate transcription by promoting the binding of positive transcription factors and blocking negative ones. (epigenie.com)
  • Methylation of lysine 4 on histone H3: intricacy of writing and reading a single epigenetic mark. (epigenie.com)
  • The mechanism of epigenetic regulation involves the CpG site methylation of promoter regions and the modification of DNA and histones by altering chromatin structure ( 9 - 11 ). (spandidos-publications.com)
  • Phosphorylation of serine residues within Histone H3 is a highly dynamic process that associates with acetylation and methylation to create combinatorial patterns which are read by specific detector proteins (A. Sawicka & C. Seiser (2012) Biochimie 94, 2193-2201). (eurogentec.com)
  • Among these post-translational modifications are different methylation states (mono-, di-, tri-methylation) on the lysine (K) residue at position 20 of histone 4 (H4K20). (uni-muenchen.de)
  • methylation of histone H4R3, as critical for developmental myelination. (hwupdate.org)
  • Decreased histone H4R3 symmetric methylation is followed by increased nuclear acetylation of H4K5, and is rescued by pharmacological inhibition of histone acetyltransferases. (hwupdate.org)
  • Together, these results identify PRMT5 as critical for oligodendrocyte differentiation and developmental myelination by modulating the cross-talk between histone arginine methylation and lysine acetylation. (hwupdate.org)
  • Its activity is thought to negatively regulate gene expression due to methylation of multiple arginine residues on nucleosomal histone tails20C22. (hwupdate.org)
  • Because the study of symmetric arginine methylation relies on the high quality of reagents, in this study we extensively characterized the specificity of all the commercially available antibodies to study this modification and selected those with the highest level of discriminatory power from other modifications (including asymmetric methylation at the same residue). (hwupdate.org)
Acetylation9
Proteins17
  • Histone H2B is one of the 5 main histone proteins involved in the structure of chromatin in eukaryotic cells. (wikipedia.org)
  • Similar to other histone proteins, histone H2B has a distinct histone fold that is optimized for histone-histone as well as histone-DNA interactions. (wikipedia.org)
  • Hyperacetylation of histone tails helps DNA-binding proteins access chromatin by weakening histone-DNA and nucleosome-nucleosome interactions. (wikipedia.org)
  • These variants, also called isoforms, are proteins that are structurally very similar to the regular histone H2B but feature some specific variations in their amino acid sequence. (wikipedia.org)
  • Histone H2B isoforms interact differently with other proteins, are found in specific regions of chromatin, have different types and numbers of post-translational modifications, and are more or less stable than regular histone H2B. (wikipedia.org)
  • Adding chemical groups to the DNA backbone and modifying histone proteins impart distinct characteristics on chromatin architecture. (frontiersin.org)
  • Recently, it was also shown to influence the stability and activity of non-histone proteins. (europa.eu)
  • In chromosomes, DNA is wrapped around proteins called histones. (news-medical.net)
  • The S phase is characterized by DNA synthesis, and in the 1960s it was discovered that histone proteins are also synthesized during this phase. (news-medical.net)
  • HDAc catalyzes the removal of acetyl groups from the lysine residues of histones and some nonhistone proteins. (medscape.com)
  • The Tudor domain binds to proteins with dimethylated arginine or lysine residues, and may also bind methylated histone tails to facilitate protein-protein interactions. (nih.gov)
  • Post-translational modifications of histone proteins play a pivotal role in DNA packaging and regulation of genome functions. (sdbonline.org)
  • Histone acetyltransferase 1 (Hat1) proteins are conserved enzymes that modify histones by acetylating lysine residues. (sdbonline.org)
  • In mice, rapamycin treatment increases expression of histone proteins and Wdfy3 transcription, and alters chromatin organisation in the small intestine, suggesting the mTORC1-histone axis is at least partially conserved in mammals and may offer new targets for anti-ageing interventions. (sdbonline.org)
  • In this study, we adopted several strategies to address this key question including: a detailed characterization of mice with cell-lineage-specific ablation of in immature oligodendrocyte progenitors or in oligodendrocytes, the use of CRISPR/Cas9 and pharmacological inhibitors to interfere with PRMT5 function in primary OPC cultures, transcriptomic analyses, and biochemical assays using synthetic proteins and modified histone peptides. (hwupdate.org)
  • We found that cells lacking 1-20 residues of H2ANtT are intolerant to copper stress, owing to the accumulation of misfolded proteins in the mutant cells. (bvsalud.org)
  • Non-histone chromosomal proteins that regulate GENE EXPRESSION through a variety of mechanisms including recognizing acetylated HISTONE LYSINE residues and through transcriptional machinery regulation. (bvsalud.org)
Lysine residues of histones1
  • Post-translational modifications of lysine residues of histones can result in a series of functional changes. (nel.edu)
Chromatin structure1
  • Histone H2B helps regulate chromatin structure and function through post-translational modifications and specialized histone variants. (wikipedia.org)
Peptides3
  • Furthermore, high-resolution X-ray crystal structures of PRDM3 and PRDM16 N-terminal peptides in complex with RBBP4 revealed binding to RBBP4 within the conserved histone H3-binding groove. (rcsb.org)
  • A polyclonal antibody targeting the dimethyl-Lysine 9 of Histone 3 was tested against various H3 peptides and shown to bind to all peptides containing the dimethyl-Lysine 9, no matter the surrounding residues. (eurogentec.com)
  • The Speedy 28-day polyclonal antibody recognizes Histone H3 K9me2 (E), independently of neighboring modifications S10ph and K14ac marks (H,I). The antibody specificity has been determined with different peptides derived from histone H3 (amino acids 1-20). (eurogentec.com)
Transcriptional2
  • It helps in deacetylation of lysine residues on the N-terminal part of the core histones (H2A, H2B, H3 and H4) and gives a tag for epigenetic repression and plays an important role in transcriptional regulation, cell cycle progression and developmental events. (novusbio.com)
  • Functions of histone H3 and H4 tail residues in stress-responsive gene transcriptional programs have been extensively studied. (bvsalud.org)
Phosphorylation2
Enzymes2
  • Covalent modifications of histones affect DNA compaction and can recruit further histone modifying enzymes. (uni-muenchen.de)
  • Mass spectrometry-based proteomics of individual ticks demonstrated persistence of mammalian host blood components, including α- and β-globin chains, histones, and mitochondrial enzymes, in Ixodes scapularis and Amblyomma americanum ticks for months after molting. (cdc.gov)
Peptide3
  • Synthesized histone H3 peptide corresponding to residues within 1-21 of human histone H3, biotin conjugated. (epigentek.com)
  • The proposal will apply an interdisciplinary approach in an international environment to maximize its chances of success: the biological role of reading domains will be evaluated with molecular biology (histone and chromatin pulldowns, ChIP-seq, confocal microscopy), biochemistry (Peptide arrays, mass spectrometry, methyltransferase activity assays) and biophysics techniques (fluorescence anisotropy, circular dichroism spectroscopy) as well as crystallography (solving the structure of reading domains). (europa.eu)
  • This peptide is Histone 3, with amino acid residues 21 to 44. (eurogentec.com)
Transcription12
  • Ubiquitinated histone H2B is often found in regions of active transcription. (wikipedia.org)
  • Specifically, the ubiquitin on histone H2B opens up and unfolds regions of chromatin allowing transcription machinery access to the promoter and coding regions of DNA. (wikipedia.org)
  • Specifically, in some lower eukaryotes, a histone H2B variant binds to a histone H2A variant called H2AZ, localizes to active genes, and supports transcription in those regions. (wikipedia.org)
  • Histone deacetylases (HDACs) are protein deacetylases that play a role in repression of gene transcription and are emerging targets in cancer therapy. (rcsb.org)
  • The histone variant mH2A1.1 interferes with transcription by down-regulating PARP-1 enzymatic activity. (nature.com)
  • The regulation of transcription of histone genes differs between eukaryotes. (news-medical.net)
  • This mechanism is hypothesized to transduce a shared signal that activates the transcription of histone genes when the G1/S phase boundary is reached. (news-medical.net)
  • A cyclin E-CDK2 substrate called NPAT has been found to be associated with histone gene clusters, and cyclin E-CDK2 bolsters activation of histone gene transcription by NPAT. (news-medical.net)
  • Therefore, NPAT could be the link between cell cycle machinery and shared heightened transcription of histone genes during S phase. (news-medical.net)
  • Histone deacetylase (HDAC) inhibitors are important epigenetic mechanisms that regulate gene expression and transcription. (pharmiweb.com)
  • Set9, a novel histone H3 methyltransferase that facilitates transcription by precluding histone tail modifications required for heterochromatin formation. (epigenie.com)
  • Histone tail residues drive many biological processes by regulating genome-wide transcription. (bvsalud.org)
Modifications3
  • Recent achievements highlight the accelerated development of epigenetics, such as the definition of a human DNA methylome at single-nucleotide resolution, the various discoveries of histone variants and modifications, the study of the CpG island in the genome, and the progress of genome-wide nucleosome positioning maps ( Baldi, 2019 ). (frontiersin.org)
  • According to the 'histone code' hypothesis, combinatorial modifications dictate a biological readout. (eurogentec.com)
  • Besides modifications of lysine residues, repressive modifications of nucleosomal histones include the symmetric dimethylation of arginines (-NG, -NG-dimethyl arginine), which is mediated by class-II protein arginine methyltransferases (PRMTs) such as PRMT513,14 and PRMT915,16. (hwupdate.org)
Specific histone1
  • Here, we hypothesized that specific histone residue(s) might be involved in facilitating gene expression such that cells mount a response to rapamycin. (researchsquare.com)
Genes4
  • The replication independent histone genes are transcribed at a relatively constant low rate, regardless of cell cycle stage. (news-medical.net)
  • However, most of the vertebrate histone genes are replication dependent and are therefore more highly expressed during the cell cycle's S phase. (news-medical.net)
  • Given that the replication dependent histone genes all activate upon entry into the S phase, it is speculated that there exists some further upstream regulatory element. (news-medical.net)
  • Their published paper is titled "Driver mutations in histone H3.3 and chromatin remodeling genes in pediatric glioblastoma. (genengnews.com)
Tails3
  • Along with a central globular domain, histone H2B has two[verification needed] flexible histone tails that extend outwards - one at the N-terminal end and one at C-terminal end. (wikipedia.org)
  • The bromodomain has been shown to bind acetylated lysine residues on histone tails. (embl-heidelberg.de)
  • Typically, RBBP4 facilitates chromatin association of the NuRD complex by binding to histone H3 tails. (rcsb.org)
Deacetylase Inhibitors1
Gene expression2
HDAc12
  • Panobinostat is a histone deacetylase (HDAc) inhibitor. (medscape.com)
  • Global Histone Deacetylase (HDAC) Inhibitors Market is Ex. (pharmiweb.com)
  • The global histone deacetylase (HDAC) inhibitors market is expected to grow from USD 97.54 million in 2018 to USD 162.76 million by 2026 at a CAGR of 7.11% during the forecast period 2019-2026. (pharmiweb.com)
  • Histone Deacetylase (HDAC) inhibitors are anti-cancer compounds that helps the body's own immune cells to recognize and attack tumour cells. (pharmiweb.com)
  • Histone Deacetylase (HDAC) inhibitors induce cancer cell cycle arrest, differentiation and cell death, reduce angiogenesis and modulate immune response. (pharmiweb.com)
  • Histone deacetylase (HDAC) inhibitors are used in psychiatry and neurology as mood stabilizers and anti-epileptics. (pharmiweb.com)
  • This has increased demand for histone deacetylase (HDAC) inhibitors that inhibit the actions of HDAC enzyme. (pharmiweb.com)
  • The global histone deacetylase (HDAC) inhibitors market is analysed on the basis of five regions namely North America, Europe, Asia Pacific, South America, and Middle East and Africa. (pharmiweb.com)
  • North America region dominated the global histone deacetylase (HDAC) inhibitors market with a market share of 38.59% in 2018. (pharmiweb.com)
  • Early adoption of innovative therapeutics, presence of sophisticated healthcare infrastructure, high R&D spending capacity, and rapid technological advancements are the factors that led to the growth of histone deacetylase (HDAC) inhibitors market in North America region. (pharmiweb.com)
  • The companies are adopting the strategies such as product innovations, mergers & acquisitions, recent developments, joint venture, collaborations, and partnership to maintain their presence in the global histone deacetylase (HDAC) inhibitors market. (pharmiweb.com)
  • A nuclear protein, HDAC-9 belongs to the histone deacetylase family with several isoforms and is inhibited by Trichostatin A (TSA) and suberoylanilide hydroxamic acid. (novusbio.com)
Isoforms4
  • While only a few isoforms of histone H2B have been studied in depth, researchers have found that histone H2B variants serve important roles. (wikipedia.org)
  • This supports the idea that isoforms of histone H2B may have specialized functions in different tissues. (wikipedia.org)
  • Many histone H2B isoforms are expressed in a DNA replication independent manner. (wikipedia.org)
  • To elucidate functional characteristics of these N-terminal residues, we compared the protein interactomes of the full-length and ΔPR isoforms of PRDM3 and its closely related paralog, PRDM16. (rcsb.org)
Acetyltransferase1
  • Structural basis for histone and phospho-histone binding by the GCN5 histone acetyltransferase. (expasy.org)
Inhibitors1
  • Chemical PARP1 inhibitors, PARP1 knockdown and mutation of ADP-ribose-binding residues in macroH2A1.1 abrogate macrodomain recruitment. (nature.com)
Deacetylases1
  • Class IIa histone deacetylases (HDACs) regulate a variety of cellular processes, including cardiac growth, bone development, and specification of skeletal muscle fiber type. (jci.org)
Repressive1
  • Ketamine treatment increased the binding of NF‑κB and permissive histone H3 lysine‑4 (H3K4)m3, but caused a decrease in the repressive histone H3K27m3 and H3K36m3 on the COX‑2 promoter ranging from ‑1,522 to ‑1,331 bp as determined by a chromatin immunoprecipitation assay. (spandidos-publications.com)
Methyltransferases1
  • KMET-READ plans to investigate the biological role of these reading domains in essential histone lysine methyltransferases - PHDs in MLL2 and MLL3 and CDs in SUV39H1 and SUV39H2. (europa.eu)
Binds2
  • To facilitate this formation, histone H2B first binds to histone H2A to form a heterodimer. (wikipedia.org)
  • We show that RBBP4 binds to the N-terminal amino acid residues of PRDM3 and PRDM16, with a dissociation constant of 3.0 μM, as measured by isothermal titration calorimetry. (rcsb.org)
Inhibition1
  • This study shows that inhibition of mTORC1 by the lifespan-extending drug rapamycin increases expression of histones H3 and H4 post-transcriptionally, through eIF3 -mediated translation. (sdbonline.org)
H3K41
  • Like all lysine residues, H3K4 can be mono, di, or tri methylated, and each have slightly different distributions. (epigenie.com)
Modification2
  • Ubiquitinase RNF20/RNF40 specifically modifies histone H2B at position K120 and this modification is need for recruitment to damaged DNA of the factors necessary for repair by the pathways of homologous recombination and non-homologous end joining. (wikipedia.org)
  • histone modification and intracellular protein breakdown. (theodora.com)
Nucleosome2
  • Two copies of histone H2B come together with two copies each of histone H2A, histone H3, and histone H4 to form the octamer core of the nucleosome to give structure to DNA. (wikipedia.org)
  • Two of these heterodimers then bind together with a heterotetramer made of histone H3 and histone H4, giving the nucleosome its characteristic disk shape. (wikipedia.org)
Mammalian1
  • The main mammalian histone subtypes ( H2A, H2B, H3, and H4 ) and the linker H1 are located in different gene clusters. (news-medical.net)
Replication independent1
  • Histones can be classified as replication dependent or replication independent, which is decided by their expression pattern during the cell cycle. (news-medical.net)
Biochemical1
  • Protein sequence information mainly consists of amino acid residue composition, biochemical features of amino acid residues and evolutionary information in terms of position-specific scoring matrices (PSSM). (nature.com)
MRNA2
  • The amount of histone mRNA increases approximately 25 to 30 times during the S phase. (news-medical.net)
  • Subsequently, partial reversibility of the ciliogenic phenotype by means of concomitant mRNA injections of the histone de-methylase Phf8, removing H4K20 mono-methyl, could be demonstrated. (uni-muenchen.de)
Orthologs1
  • The sequence, structure, and habitat of selected histone-like nucleoid-structuring (H-NS) orthologs. (elifesciences.org)
Polyclonal1
  • The polyclonal anti-H3 K9me2 is able to recognize Histone H3 multi-modified on K9me2 S10ph K14ac. (eurogentec.com)
Amino acids2
Specificity1
  • The knowledge about DNA-binding residues, binding specificity and binding affinity helps to not only understand the recognition mechanism of protein-DNA complex, but also give clues for protein function annotation. (nature.com)
Ubiquitination1
Variant3
  • Notably, histone macroH2A1.1 senses PARP1 activation, transiently compacts chromatin, reduces the recruitment of DNA damage factor Ku70-Ku80 and alters γ-H2AX patterns, whereas the splice variant macroH2A1.2, which is deficient in poly-ADP-ribose binding, does not mediate chromatin rearrangements upon PARP1 activation. (nature.com)
  • Poly(ADP-ribose) polymerase 1 is inhibited by a histone H2A variant, MacroH2A, and contributes to silencing of the inactive X chromosome. (nature.com)
  • Hat1 function and the presence of these acetyl marks are not required for the nuclear transport of histone H4 as histone variant His4r retained its nuclear localization both in Hat1 mutants and in His4r-K5R-K12R double point mutants. (sdbonline.org)
Mutations3
  • Bullock and Fersht 8 have shown that mutations of DNA-binding residues, such as those on the tumor repressor protein P53, may predispose individuals to cancer. (nature.com)
  • Independent researchers have identified mutations in the histone H3.3-coding gene H3F3A in a large proportion of pediatric cases of gliobastoma multiforme (GBM) and diffuse intrinsic pontine glioma (DIPG), an aggressive brainstem astrocytic tumor that occurs almost exclusively in children. (genengnews.com)
  • Both teams suggest their respective works are the first to identify somatic mutations in histone H3. (genengnews.com)
Chromosome1
  • Kornberg 7 has presented that the interactions between DNA and histones are involved in chromosome packaging in the cell nucleus. (nature.com)
Structural2
  • Histone H2B is a structural protein that helps organize eukaryotic DNA. (wikipedia.org)
  • The family is defined by structural similarity of their kinase domains and their capability to autophosphorylate on tyrosine residues. (antibodies-online.com)
Variants1
  • There are sixteen variants of histone H2B found in humans, thirteen of which are expressed in regular body cells and three of which are only expressed in the testes. (wikipedia.org)
Distinct1
  • Here we show that distinct macrodomains, including those of histone macroH2A1.1, are recruited to sites of PARP1 activation induced by laser-generated DNA damage. (nature.com)
Recognize1
  • Double chromodomains cooperate to recognize the methylated histone H3 tail. (epigenie.com)
Alterations1
  • The St. Jude-Washington University team reports its findings in Nature Genetics in a paper titled "Somatic histone H3 alterations in pediatric diffuse intrinsic pontine gliomas and nonbrainstem glioblastomas. (genengnews.com)
Interactions1
  • Nonpolar and ionic interactions in histone-like nucleoid-structuring (H-NS) site1 and site2 dimers. (elifesciences.org)
Rapamycin4
  • Herein, we screened a histone mutant collection and report that one of the mutants, H2B R95A, is strikingly resistant to rapamycin due to a defective cell cycle arrest. (researchsquare.com)
  • Indeed, it has been shown that a histone mutant H3 K56A causes yeast cells to display sensitivity to rapamycin, however, this mutation also sensitizes cells to a variety of DNA damaging agents, suggesting that the H3 K56A mutation has a broader role rather than a specific function towards rapamycin stress [22, 23]. (researchsquare.com)
  • We, therefore, searched a collection of 442 histone mutants [24] for ones that failed to respond to rapamycin in an attempt to identify molecular pathways critical for rapamycin response at the level of chromatin. (researchsquare.com)
  • We report the identification of nine histone mutants, eight of which showed varying resistance to rapamycin, as compared to the other mutants and the wild type (WT). (researchsquare.com)
Drosophila1
Asymmetric1
  • Biased fork movement coupled with a strand preference in histone incorporation would explain how asymmetric old and new H3 and H4 are established during replication. (sdbonline.org)
Molecular1
  • Molecular basis for site-specific read-out of histone H3K4me3 by the BPTF PHD finger of NURF. (epigenie.com)
Activates1
  • Hypoxia changes the cellular redox state and activates class III histone deacetylase sirtuin1 (SIRT1). (molvis.org)
Localization1
  • This is a key paper defining the localization of all common histone methylations in the human genome. (epigenie.com)