Epigenetic Regulation of Learning and Memory by Drosophila EHMT/G9a
Publication Date
January 04, 2011
Journal
PLOS Biology
Authors
Jamie M. Kramer, Korinna Kochinke, Merel A. W. Oortveld, Hendrik Marks, et al
Volume
9
Issue
1
Pages
e1000569
DOI
https://dx.plos.org/10.1371/journal.pbio.1000569
Publisher URL
http://journals.plos.org/plosbiology/article?id=10.1371%2Fjournal.pbio.1000569
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/21245904
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3014924
Europe PMC
http://europepmc.org/abstract/MED/21245904
Web of Science
000286595000012
Scopus
79851484936
Mendeley
http://www.mendeley.com/research/epigenetic-regulation-learning-memory-drosophila-ehmtg9a
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{"title"=>"Epigenetic regulation of learning and memory by Drosophila EHMT/G9a", "type"=>"journal", "authors"=>[{"first_name"=>"Jamie M.", "last_name"=>"Kramer", "scopus_author_id"=>"7402417460"}, {"first_name"=>"Korinna", "last_name"=>"Kochinke", "scopus_author_id"=>"36959413800"}, {"first_name"=>"Merel A.W.", "last_name"=>"Oortveld", "scopus_author_id"=>"35345521100"}, {"first_name"=>"Hendrik", "last_name"=>"Marks", "scopus_author_id"=>"15081168100"}, {"first_name"=>"Daniela", "last_name"=>"Kramer", "scopus_author_id"=>"57200844768"}, {"first_name"=>"Eiko K.", "last_name"=>"de Jong", "scopus_author_id"=>"57197122401"}, {"first_name"=>"Zoltan", "last_name"=>"Asztalos", "scopus_author_id"=>"6602664943"}, {"first_name"=>"J. Timothy", "last_name"=>"Westwood", "scopus_author_id"=>"7004670277"}, {"first_name"=>"Hendrik G.", "last_name"=>"Stunnenberg", "scopus_author_id"=>"7006024572"}, {"first_name"=>"Marla B.", "last_name"=>"Sokolowski", "scopus_author_id"=>"7103249605"}, {"first_name"=>"Krystyna", "last_name"=>"Keleman", "scopus_author_id"=>"6602872492"}, {"first_name"=>"Huiqing", "last_name"=>"Zhou", "scopus_author_id"=>"12792008500"}, {"first_name"=>"Hans", "last_name"=>"van Bokhoven", "scopus_author_id"=>"7005587499"}, {"first_name"=>"Annette", "last_name"=>"Schenck", "scopus_author_id"=>"7006174008"}], "year"=>2011, "source"=>"PLoS Biology", "identifiers"=>{"issn"=>"15449173", "scopus"=>"2-s2.0-79851484936", "sgr"=>"79851484936", "pui"=>"361245359", "isbn"=>"1545-7885 (Electronic)\\r1544-9173 (Linking)", "pmid"=>"21245904", "doi"=>"10.1371/journal.pbio.1000569"}, "id"=>"e37d099e-2a4b-33ca-a3ff-4e49dc1a620e", "abstract"=>"The epigenetic modification of chromatin structure and its effect on complex neuronal processes like learning and memory is an emerging field in neuroscience. However, little is known about the \"writers\" of the neuronal epigenome and how they lay down the basis for proper cognition. Here, we have dissected the neuronal function of the Drosophila euchromatin histone methyltransferase (EHMT), a member of a conserved protein family that methylates histone 3 at lysine 9 (H3K9). EHMT is widely expressed in the nervous system and other tissues, yet EHMT mutant flies are viable. Neurodevelopmental and behavioral analyses identified EHMT as a regulator of peripheral dendrite development, larval locomotor behavior, non-associative learning, and courtship memory. The requirement for EHMT in memory was mapped to 7B-Gal4 positive cells, which are, in adult brains, predominantly mushroom body neurons. Moreover, memory was restored by EHMT re-expression during adulthood, indicating that cognitive defects are reversible in EHMT mutants. To uncover the underlying molecular mechanisms, we generated genome-wide H3K9 dimethylation profiles by ChIP-seq. Loss of H3K9 dimethylation in EHMT mutants occurs at 5% of the euchromatic genome and is enriched at the 5' and 3' ends of distinct classes of genes that control neuronal and behavioral processes that are corrupted in EHMT mutants. Our study identifies Drosophila EHMT as a key regulator of cognition that orchestrates an epigenetic program featuring classic learning and memory genes. Our findings are relevant to the pathophysiological mechanisms underlying Kleefstra Syndrome, a severe form of intellectual disability caused by mutations in human EHMT1, and have potential therapeutic implications. Our work thus provides novel insights into the epigenetic control of cognition in health and disease.", "link"=>"http://www.mendeley.com/research/epigenetic-regulation-learning-memory-drosophila-ehmtg9a", "reader_count"=>244, "reader_count_by_academic_status"=>{"Unspecified"=>4, "Professor > Associate Professor"=>18, "Librarian"=>2, "Student > Doctoral Student"=>8, "Researcher"=>58, "Student > Ph. D. Student"=>67, "Student > Postgraduate"=>5, "Student > Master"=>32, "Other"=>8, "Student > Bachelor"=>29, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>10}, "reader_count_by_user_role"=>{"Unspecified"=>4, "Professor > Associate Professor"=>18, "Librarian"=>2, "Student > Doctoral Student"=>8, "Researcher"=>58, "Student > Ph. D. 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Scopus | Further Information

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  • {"files"=>["https://ndownloader.figshare.com/files/808464"], "description"=>"<p>(a) Larval crawling paths from <i>EHMT</i><sup>+</sup>, <i>EHMT<sup>DD1</sup></i>, and <i>EHMT<sup>DD2</sup></i>. (b–c) Quantification of larval locomotory patterns revealed no significant difference in the (b) total path length but showed a significant increase in (c) side track length in both <i>EHMT<sup>DD1</sup></i> (red bars) and <i>EHMT<sup>DD2</sup></i> (orange bars). Data are normalized to EHMT<sup>+</sup> and error bars represent the standard error of the mean. <i>N</i> = 149, 148, and 149 for <i>EHMT</i><sup>+</sup>, <i>EHMT<sup>DD1</sup></i>, and <i>EHMT<sup>DD2</sup></i>, respectively. (*) indicates a significant difference to <i>EHMT</i><sup>+</sup> (<i>p</i><0.05, Kruskal-Wallis and Mann-Whitney tests).</p>", "links"=>[], "tags"=>["mutants", "altered", "larval", "locomotory"], "article_id"=>478817, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g004", "stats"=>{"downloads"=>2, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_EHMT_mutants_show_altered_larval_locomotory_behavior_/478817", "title"=>"EHMT mutants show altered larval locomotory behavior.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:26:57"}
  • {"files"=>["https://ndownloader.figshare.com/files/808762"], "description"=>"<p>(a) Venn diagram showing the overlap in genes associated with either Upstream LOMBs or Downstream LOMBs. (b) Heat map showing the significance of enrichment or depletion for gene ontology terms describing biological processes associated with: genes containing LOMBs (All LOMBs—5,136); genes containing LOMBs within 1 kb upstream of the transcriptional start site (Upstream LOMBs—1,229); genes containing LOMBs within 1 kb downstream of the polyA site (Downstream LOMBs—1,712); and genes that do not contain any LOMBs (No LOMBs—10,238). Complete lists of gene IDs for these groups are provided in <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000569#pbio.1000569.s010\" target=\"_blank\">Table S3</a>. Color scale indicates the significance of enrichment as determined using GOToolBox to perform (<a href=\"http://genome.crg.es/GOToolBox\" target=\"_blank\">http://genome.crg.es/GOToolBox</a>) a hypergeometric test with Benjamini & Hochberg correction. Darkest colors indicate <i>p</i> values ≤10<sup>−7</sup>. <i>ns</i>  =  non-significant (<i>p</i>>0.05).</p>", "links"=>[], "tags"=>["ehmt"], "article_id"=>479120, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g007", "stats"=>{"downloads"=>2, "page_views"=>19, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Bioinformatic_analysis_of_EHMT_target_genes_/479120", "title"=>"Bioinformatic analysis of EHMT target genes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:32:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/808240"], "description"=>"<p>(a) Schematic depiction of the genomic region containing <i>EHMT</i> and the location of the <i>EHMT</i> deletions. Blue boxes indicate <i>EHMT</i> exons. The red bar shows the location of the predicted translational start site. The black bar and triangle indicate the location of the P-element insertion <i>KG01242</i>. The green bar marks the epitope of the EHMT antibody. The size and location of the <i>EHMT</i> deletions <i>DD1</i> and <i>DD2</i> (downstream deletions 1 and 2) is indicated. (b) Western blot using anti-EHMT and embryonic protein extracts from <i>EHMT</i><sup>+</sup> (lane 1), <i>EHMT<sup>DD1</sup></i> (lane 2), and <i>EHMT<sup>DD2</sup></i> (lane 3).The EHMT band (arrowhead) is positioned in accordance with a predicted size of 180 kDa. This band is absent in the deletion lines and no extra bands were observed. α-tubulin was used as loading control. (c) <i>EHMT</i><sup>+</sup>, <i>EHMT<sup>DD1</sup></i>, and <i>EHMT<sup>DD2</sup></i> embryos at blastoderm stage, stained with anti-EHMT (green) and DAPI (magenta). EHMT staining is nuclear in wild-type and absent in mutant <i>EHMT</i> embryos. Scale bar is 100 µm.</p>", "links"=>[], "tags"=>["characterization"], "article_id"=>478595, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g002", "stats"=>{"downloads"=>3, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Molecular_characterization_of_EHMT_alleles_/478595", "title"=>"Molecular characterization of <i>EHMT</i> alleles.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:23:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/403508", "https://ndownloader.figshare.com/files/403528", "https://ndownloader.figshare.com/files/403540", "https://ndownloader.figshare.com/files/403553", "https://ndownloader.figshare.com/files/403570", "https://ndownloader.figshare.com/files/403595", "https://ndownloader.figshare.com/files/403623", "https://ndownloader.figshare.com/files/403641", "https://ndownloader.figshare.com/files/403659", "https://ndownloader.figshare.com/files/403682", "https://ndownloader.figshare.com/files/403696", "https://ndownloader.figshare.com/files/403729"], "description"=>"<div><p>The epigenetic modification of chromatin structure and its effect on complex neuronal processes like learning and memory is an emerging field in neuroscience. However, little is known about the “writers” of the neuronal epigenome and how they lay down the basis for proper cognition. Here, we have dissected the neuronal function of the <em>Drosophila</em> euchromatin histone methyltransferase (EHMT), a member of a conserved protein family that methylates histone 3 at lysine 9 (H3K9). EHMT is widely expressed in the nervous system and other tissues, yet EHMT mutant flies are viable. Neurodevelopmental and behavioral analyses identified EHMT as a regulator of peripheral dendrite development, larval locomotor behavior, non-associative learning, and courtship memory. The requirement for EHMT in memory was mapped to 7B-Gal4 positive cells, which are, in adult brains, predominantly mushroom body neurons. Moreover, memory was restored by EHMT re-expression during adulthood, indicating that cognitive defects are reversible in EHMT mutants. To uncover the underlying molecular mechanisms, we generated genome-wide H3K9 dimethylation profiles by ChIP-seq. Loss of H3K9 dimethylation in EHMT mutants occurs at 5% of the euchromatic genome and is enriched at the 5′ and 3′ ends of distinct classes of genes that control neuronal and behavioral processes that are corrupted in EHMT mutants. Our study identifies <em>Drosophila</em> EHMT as a key regulator of cognition that orchestrates an epigenetic program featuring classic learning and memory genes. Our findings are relevant to the pathophysiological mechanisms underlying Kleefstra Syndrome, a severe form of intellectual disability caused by mutations in human <em>EHMT1</em>, and have potential therapeutic implications. Our work thus provides novel insights into the epigenetic control of cognition in health and disease.</p></div>", "links"=>[], "tags"=>["epigenetic"], "article_id"=>139720, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>["https://dx.doi.org/10.1371/journal.pbio.1000569.s001", "https://dx.doi.org/10.1371/journal.pbio.1000569.s002", "https://dx.doi.org/10.1371/journal.pbio.1000569.s003", "https://dx.doi.org/10.1371/journal.pbio.1000569.s004", "https://dx.doi.org/10.1371/journal.pbio.1000569.s005", "https://dx.doi.org/10.1371/journal.pbio.1000569.s006", "https://dx.doi.org/10.1371/journal.pbio.1000569.s007", "https://dx.doi.org/10.1371/journal.pbio.1000569.s008", "https://dx.doi.org/10.1371/journal.pbio.1000569.s009", "https://dx.doi.org/10.1371/journal.pbio.1000569.s010", "https://dx.doi.org/10.1371/journal.pbio.1000569.s011", "https://dx.doi.org/10.1371/journal.pbio.1000569.s012"], "stats"=>{"downloads"=>34, "page_views"=>16, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Epigenetic_Regulation_of_Learning_and_Memory_by_Drosophila_EHMT_G9a/139720", "title"=>"Epigenetic Regulation of Learning and Memory by <em>Drosophila</em> EHMT/G9a", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2011-01-04 02:42:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/808539"], "description"=>"<p>(a–c) Jump reflex habituation. (a–b) Habituation was measured in (a) male flies of the genotypes <i>EHMT<sup>+</sup>/Y</i> (black diamonds), <i>EHMT<sup>DD1</sup>/Y</i> (red squares), and <i>EHMT<sup>DD2</sup>/Y</i> (orange triangles) and (b) female flies of the genotypes <i>EHMT<sup>+</sup>/EHMT<sup>+</sup></i> (black diamonds) and <i>EHMT<sup>DD1</sup>/EHMT<sup>DD2</sup></i> (red circles). Jumping was induced by repeated light-off pulses for 100 trials. (c) The mean number of trials to criterion was significantly higher for <i>EHMT</i> mutants (orange and red bars) than for <i>EHMT<sup>+</sup></i> flies (black bars) after training with a 1 s inter-trial interval. (*) indicates a significant difference (<i>p</i><0.001, one-way ANOVA and Bonferroni tests). (d–f) Learning and memory in the courtship conditioning paradigm. (d) The Learning Index (LI) of <i>EHMT<sup>DD1</sup></i> males was normal at 0 min after training but was significantly reduced at 30 min after training (short term memory—STM) and 24 h after training (long term memory—LTM). (e) The LI of <i>EHMT<sup>DD2</sup></i> males was also affected in short term memory and was rescued by expression of <i>EHMT</i> with the pan neuronal <i>elav-Gal4</i> driver and the <i>7B-Gal4</i> driver, which is predominantly expressed in the mushroom bodies in adult fly brains (orange bars). In the <i>EHMT<sup>+</sup></i> background, expression of EHMT with <i>elav-Gal4</i> had no adverse effect, while expression with <i>7B-Gal4</i> caused a significantly decreased LI (black bars). (f) Short term memory was rescued in the <i>EHMT<sup>DD1</sup></i> mutant background by induced expression of EHMT during adulthood, using the <i>hsGal4</i> driver. See Experimental Procedures for details on heat-shock conditions and courtship training. Error bars represent standard error of the mean. (*) indicates a significant difference (Kruskall-Wallis and Mann-Whitney tests).</p>", "links"=>[], "tags"=>["affects", "non-associative", "courtship"], "article_id"=>478891, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g005", "stats"=>{"downloads"=>3, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_EHMT_affects_non_associative_learning_and_courtship_memory_/478891", "title"=>"EHMT affects non-associative learning and courtship memory.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:28:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/808093"], "description"=>"<p>(a) A phylogenetic tree generated using the neighbor joining method showing the evolutionary relationship between <i>Drosophila</i> EHMT and the mouse and human orthologs of EHMT1 and EHMT2/G9a. Analysis was performed with the Vector NTI software (Invitrogen). The scale bar indicates phylogenetic distance. (b) Frontal view of an adult <i>Drosophila</i> brain stained with an antibody to EHMT (green) and counterstained with DAPI (magenta). EHMT is widely expressed but excluded from neuropilar regions such as the mushroom body calyx (asterisk in merge panel). (c–d) Horizontal view of a third instar larval ventral nerve cord (vnc) stained with an anti-EHMT antibody (green) and counterstained with (c) anti-elav (magenta) to highlight neuronal nuclei or with (d) anti-repo (magenta) labeling glial nuclei. EHMT colocalizes with elav and repo but appears to be expressed at a lower level in glia. (e) The ventral cluster of multiple dendrite neurons of the larval body wall labeled with memGFP (green) using the <i>109(2)80-Gal4</i> driver and stained with anti-EHMT (red) and DAPI (blue). Arrowheads point to the midline of the brain (in b) and vnc (in c, d). Scale bars represent 100 µm in (b), 50 µm in (c–d), and 10 µm in (e). Anterior is to the left (c–e).</p>", "links"=>[], "tags"=>["phylogeny"], "article_id"=>478448, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g001", "stats"=>{"downloads"=>4, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_EHMT_phylogeny_and_expression_in_the_nervous_system_/478448", "title"=>"<i>EHMT</i> phylogeny and expression in the nervous system.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:20:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/808648"], "description"=>"<p>(a) Histogram showing the distribution of H3K9me2 methylation ratios in logarithmic scale (log # of sequenced tags in wt/# of sequenced tags in mt) after binning of the euchromatic genome into 384,944 300-bp segments. The mean (0.2±1) and 2× the standard deviation is indicated. The region outside +2× the standard deviation is comprised of 19,258 bins, in which methylation is greater than 4.6-fold higher in <i>EHMT<sup>+</sup></i> than in <i>EHMT<sup>DD1</sup></i>. These are referred to as loss of methylation bins (LOMBs). In contrast, the region outside −2× standard deviation contains only 50 bins. (b) Two examples of genomic loci with loss of methylation (LOM) in <i>EHMT</i> mutants. The <i>orb2</i> genes has a downstream LOM region (top panel) while the <i>Sod</i> gene has an upstream LOM region. Scale is indicated. (c) The positional distribution of all bins in relation to genes is compared to the distribution of LOMBs. These distributions are significantly different (<i>p</i><0.001; chi-squared test). LOMBs are significantly enriched within 1 kb upstream of the transcriptional start site (tss) and 1 kb downstream of the polyadenylation site (polyA), by 1.6- and 3.3-fold, respectively (<i>p</i><0.001, hypergeometric test with Bonferroni correction). (d–e) Average number of sequenced tags in <i>EHMT</i><sup>+</sup> (black) and <i>EHMT<sup>DD1</sup></i>(orange) upstream of the tss (d) and near the polyA site (e) for all genes (left panels); for genes that are >2.5-fold downregulated in <i>EHMT</i> mutants (middle panels); and for genes that are >2.5-fold upregulated in <i>EHMT</i> mutants (right panels).</p>", "links"=>[], "tags"=>["methylates", "discrete", "regions", "euchromatic"], "article_id"=>478999, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g006", "stats"=>{"downloads"=>4, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_EHMT_methylates_discrete_regions_of_the_euchromatic_genome_/478999", "title"=>"EHMT methylates discrete regions of the euchromatic genome.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:29:59"}
  • {"files"=>["https://ndownloader.figshare.com/files/808358"], "description"=>"<p>(a, b) Representative confocal images of ventral type 4 md neurons from (a) <i>EHMT</i><sup>+</sup> and (b) <i>EHMT<sup>DD2</sup></i> third instar larvae labeled by expression of <i>UAS-memGFP</i> using the <i>477-Gal4</i> driver. I, II, III, and IV label the primary dendrite branches, which are conserved in all neurons examined. Scale bars represent 50 µm. (a', b') High magnifications of neurons shown in (a) and (b), respectively. Scale bars represent 10 µm. (c) Quantitative evaluation of dendrite ends per field of view, as shown in (a–b). Note that dendrite branching was significantly reduced in <i>EHMT</i> mutants. 477<i>-Gal4</i> driven expression of <i>UAS-EHMT</i> caused a non-significant reduction of dendrite branches in the <i>EHMT</i><sup>+</sup> background (black bars) and restored the number of dendrite ends in <i>EHMT<sup>DD1</sup></i> (red bars) and <i>EHMT<sup>DD2</sup></i> (orange bars). Error bars represent standard error of the mean. Complete genotypes are indicated. Sample sizes were <i>n</i> = 19, 20, 18 for the mutant conditions and 9, 8, and 11 for the rescue experiment, respectively, in <i>EHMT</i><sup>+</sup>, <i>EHMT<sup>DD1</sup></i>, and <i>EHMT<sup>DD2</sup></i>. (*) represents a significant difference to <i>EHMT<sup>+</sup>; 477-Gal4, UAS-memGFP/+</i> (<i>p</i><0.05, one-way ANOVA and Bonferroni tests).</p>", "links"=>[], "tags"=>["regulates", "dendrite"], "article_id"=>478717, "categories"=>["Neuroscience", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Jamie M. Kramer", "Korinna Kochinke", "Merel A. W. Oortveld", "Hendrik Marks", "Daniela Kramer", "Eiko K. de Jong", "Zoltan Asztalos", "J. Timothy Westwood", "Hendrik G. Stunnenberg", "Marla B. Sokolowski", "Krystyna Keleman", "Huiqing Zhou", "Hans van Bokhoven", "Annette Schenck"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000569.g003", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_EHMT_regulates_dendrite_branching_/478717", "title"=>"<i>EHMT</i> regulates dendrite branching.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-01-04 02:25:17"}

PMC Usage Stats | Further Information

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  • {"unique-ip"=>"19", "full-text"=>"20", "pdf"=>"7", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}
  • {"unique-ip"=>"22", "full-text"=>"25", "pdf"=>"9", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"16", "full-text"=>"20", "pdf"=>"7", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
  • {"unique-ip"=>"71", "full-text"=>"35", "pdf"=>"8", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"17", "supp-data"=>"20", "cited-by"=>"1", "year"=>"2019", "month"=>"10"}
  • {"unique-ip"=>"23", "full-text"=>"22", "pdf"=>"8", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"14", "cited-by"=>"0", "year"=>"2019", "month"=>"12"}
  • {"unique-ip"=>"25", "full-text"=>"25", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"14", "cited-by"=>"0", "year"=>"2020", "month"=>"2"}

Relative Metric

{"start_date"=>"2011-01-01T00:00:00Z", "end_date"=>"2011-12-31T00:00:00Z", "subject_areas"=>[{"subject_area"=>"/Biology and life sciences/Cell biology", "average_usage"=>[289, 559, 699, 817, 931, 1031, 1127, 1214, 1297, 1370, 1444, 1515, 1584, 1656, 1726, 1797, 1862, 1930, 1996, 2061, 2125, 2190, 2250, 2311, 2373, 2435, 2496, 2563, 2630, 2692, 2760, 2824, 2898, 2960, 3019, 3089, 3143]}, {"subject_area"=>"/Biology and life sciences/Neuroscience", "average_usage"=>[306, 540, 664, 768, 864, 959, 1036, 1121, 1199, 1277, 1341, 1411, 1477, 1543, 1609, 1675, 1739, 1802, 1868, 1933, 1995, 2050, 2107, 2164, 2220, 2277, 2327, 2374, 2436, 2488, 2542, 2594, 2650, 2705, 2769, 2817, 2859]}, {"subject_area"=>"/Biology and life sciences/Psychology", "average_usage"=>[307, 557, 674, 784, 873, 964, 1048, 1134, 1206, 1285, 1350, 1417, 1486, 1553, 1610, 1683, 1752, 1824, 1890, 1953, 2003, 2067, 2132, 2198, 2255, 2304, 2353, 2414, 2468, 2512, 2576, 2625, 2687, 2750, 2808, 2849, 2888]}, {"subject_area"=>"/Social sciences", "average_usage"=>[340, 584, 708, 809, 915, 1005, 1089, 1174, 1254, 1334, 1408, 1478, 1563, 1648, 1716, 1791, 1847, 1915, 1979, 2044, 2105, 2163, 2218, 2297, 2382, 2436, 2493, 2553, 2618, 2682, 2741, 2786, 2847, 2937, 2989, 3039, 3102]}, {"subject_area"=>"/Social sciences/Psychology", "average_usage"=>[307, 557, 674, 784, 873, 964, 1048, 1134, 1206, 1285, 1350, 1417, 1486, 1553, 1610, 1683, 1752, 1824, 1890, 1953, 2003, 2067, 2132, 2198, 2255, 2304, 2353, 2414, 2468, 2512, 2576, 2625, 2687, 2750, 2808, 2849, 2888]}]}
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