Non-Coding Changes Cause Sex-Specific Wing Size Differences between Closely Related Species of Nasonia
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{"title"=>"Non-coding changes cause sex-specific wing size differences between closely related species of Nasonia", "type"=>"journal", "authors"=>[{"first_name"=>"David W.", "last_name"=>"Loehlin", "scopus_author_id"=>"35191956800"}, {"first_name"=>"Deodoro C.S.G.", "last_name"=>"Oliveira", "scopus_author_id"=>"35076438100"}, {"first_name"=>"Rachel", "last_name"=>"Edwards", "scopus_author_id"=>"35733746300"}, {"first_name"=>"Jonathan D.", "last_name"=>"Giebel", "scopus_author_id"=>"21933513400"}, {"first_name"=>"Michael E.", "last_name"=>"Clark", "scopus_author_id"=>"7404528050"}, {"first_name"=>"M. Victoria", "last_name"=>"Cattani", "scopus_author_id"=>"27267581800"}, {"first_name"=>"Louis", "last_name"=>"Van De Zande", "scopus_author_id"=>"6701442824"}, {"first_name"=>"Eveline C.", "last_name"=>"Verhulst", "scopus_author_id"=>"35076721400"}, {"first_name"=>"Leo W.", "last_name"=>"Beukeboom", "scopus_author_id"=>"7003336203"}, {"first_name"=>"Monica", "last_name"=>"Muñoz-Torres", "scopus_author_id"=>"21934921800"}, {"first_name"=>"John H.", "last_name"=>"Werren", "scopus_author_id"=>"7006624294"}], "year"=>2010, "source"=>"PLoS Genetics", "identifiers"=>{"pmid"=>"20090834", "doi"=>"10.1371/journal.pgen.1000821", "sgr"=>"76749129414", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)", "scopus"=>"2-s2.0-76749129414", "issn"=>"15537390", "pui"=>"358308133"}, "id"=>"d540c653-c50b-3a91-9499-098b968d9ba8", "abstract"=>"The genetic basis of morphological differences among species is still poorly understood. We investigated the genetic basis of sex-specific differences in wing size between two closely related species of Nasonia by positional cloning a major male-specific locus, wing-size1 (ws1). Male wing size increases by 45% through cell size and cell number changes when the ws1 allele from N. giraulti is backcrossed into a N. vitripennis genetic background. A positional cloning approach was used to fine-scale map the ws1 locus to a 13.5 kilobase region. This region falls between prospero (a transcription factor involved in neurogenesis) and the master sex-determining gene doublesex. It contains the 5'-UTR and cis-regulatory domain of doublesex, and no coding sequence. Wing size reduction correlates with an increase in doublesex expression level that is specific to developing male wings. Our results indicate that non-coding changes are responsible for recent divergence in sex-specific morphology between two closely related species. We have not yet resolved whether wing size evolution at the ws1 locus is caused by regulatory alterations of dsx or prospero, or by another mechanism. This study demonstrates the feasibility of efficient positional cloning of quantitative trait loci (QTL) involved in a broad array of phenotypic differences among Nasonia species.", "link"=>"http://www.mendeley.com/research/noncoding-changes-cause-sexspecific-wing-size-differences-between-closely-related-species-nasonia", "reader_count"=>72, "reader_count_by_academic_status"=>{"Unspecified"=>5, "Professor > Associate Professor"=>6, "Researcher"=>16, "Student > Doctoral Student"=>1, "Student > Ph. D. Student"=>24, "Student > Postgraduate"=>2, "Other"=>3, "Student > Master"=>5, "Student > Bachelor"=>7, "Professor"=>3}, "reader_count_by_user_role"=>{"Unspecified"=>5, "Professor > Associate Professor"=>6, "Researcher"=>16, "Student > Doctoral Student"=>1, "Student > Ph. D. Student"=>24, "Student > Postgraduate"=>2, "Other"=>3, "Student > Master"=>5, "Student > Bachelor"=>7, "Professor"=>3}, "reader_count_by_subject_area"=>{"Unspecified"=>5, "Environmental Science"=>1, "Biochemistry, Genetics and Molecular Biology"=>9, "Agricultural and Biological Sciences"=>56, "Neuroscience"=>1}, "reader_count_by_subdiscipline"=>{"Neuroscience"=>{"Neuroscience"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>56}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>9}, "Unspecified"=>{"Unspecified"=>5}, "Environmental Science"=>{"Environmental Science"=>1}}, "reader_count_by_country"=>{"Netherlands"=>3, "Argentina"=>1, "United States"=>3, "Japan"=>1, "United Kingdom"=>1, "Germany"=>2}, "group_count"=>3}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/867449"], "description"=>"<p>(A) Recombination map of <i>ws1</i> and flanking phenotypic markers used for positional cloning. (B) Genome map of the region around <i>ws1</i> including gene annotations. (C) Genotype and phenotype of recombinants near <i>ws1</i>. Black: <i>N. giraulti,</i> White: <i>N. vitripennis</i>. (D) Map of sequence features in the 13.5kb <i>ws1</i> locus. Triangles: Insertions, with size given in base pairs (bp). Letters: Microsatellite repeats, with size given in bp. Blue: <i>dsx</i> 5′-UTR.</p>", "links"=>[], "tags"=>["maps"], "article_id"=>537901, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.g003", "stats"=>{"downloads"=>0, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Positional_cloning_ws1_maps_to_the_doublesex_locus_/537901", "title"=>"Positional cloning: <i>ws1</i> maps to the <i>doublesex</i> locus.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-01-15 02:11:41"}
  • {"files"=>["https://ndownloader.figshare.com/files/867559"], "description"=>"<p>Expression level change was estimated by quantitative RT–PCR. Note that the <i>vitripennis dsx</i> protein-coding region is present in both genotypes (i.e., the <i>giraulti</i> region of <i>ws1<sub>g</sub>V_40kb</i> does not include the <i>dsx</i> coding sequence). Mean expression ratios ± standard errors are shown. Expression ratios greater than 1 indicate higher male <i>dsx</i> (<i>dsx<sup>M</sup></i>) transcript level in <i>ws1<sub>v</sub>V</i> than in <i>ws1<sub>g</sub>V_40kb</i>. Sample size indicates number of independent biological replicates.</p>", "links"=>[], "tags"=>["developmental biology/developmental evolution", "evolutionary biology/developmental evolution", "evolutionary biology/evolutionary and comparative genetics", "evolutionary biology/morphogenesis and cell biology", "genetics and genomics/animal genetics", "genetics and genomics/complex traits", "genetics and genomics/gene discovery"], "article_id"=>537996, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.g004", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Change_in_doublesex_expression_due_to_ws1_/537996", "title"=>"Change in <i>doublesex</i> expression due to <i>ws1.</i>", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-01-15 02:13:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/867353"], "description"=>"<p>The diagram illustrates a screen for fine-scale recombinants (marked with *) between <i>ws1<sub>g</sub></i> and a lethal located 0.6cM away. Because of the lethal (<i>l</i>), the only live haploid males with <i>ws1<sub>g</sub></i> (large wings, <i>w</i>) are recombinant. The <i>blue</i> (<i>b</i>) marker is used as a second phenotypic marker and as a way to identify recombinants on the other side of <i>ws1.</i> Gray bar: introgression (<i>N. giraulti</i>) sequence. White bar: <i>N. vitripennis</i> sequence. Proportions shown are based on estimates of recombination rates between the markers (<a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1000821#pgen-1000821-g003\" target=\"_blank\">Figure 3</a>) and assuming no double recombinants.</p>", "links"=>[], "tags"=>["cloning", "linked"], "article_id"=>537799, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.g002", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Positional_cloning_in_Nasonia_using_linked_lethals_/537799", "title"=>"Positional cloning in <i>Nasonia</i> using linked lethals.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-01-15 02:09:59"}
  • {"files"=>["https://ndownloader.figshare.com/files/430692", "https://ndownloader.figshare.com/files/430717", "https://ndownloader.figshare.com/files/430747", "https://ndownloader.figshare.com/files/430771"], "description"=>"<div><p>The genetic basis of morphological differences among species is still poorly understood. We investigated the genetic basis of sex-specific differences in wing size between two closely related species of <em>Nasonia</em> by positional cloning a major male-specific locus, <em>wing-size1</em> (<em>ws1</em>). Male wing size increases by 45% through cell size and cell number changes when the <em>ws1</em> allele from <em>N. giraulti</em> is backcrossed into a <em>N. vitripennis</em> genetic background. A positional cloning approach was used to fine-scale map the <em>ws1</em> locus to a 13.5 kilobase region. This region falls between <em>prospero</em> (a transcription factor involved in neurogenesis) and the master sex-determining gene <em>doublesex</em>. It contains the 5′-UTR and <em>cis-</em>regulatory domain of <em>doublesex</em>, and no coding sequence. Wing size reduction correlates with an increase in <em>doublesex</em> expression level that is specific to developing male wings. Our results indicate that non-coding changes are responsible for recent divergence in sex-specific morphology between two closely related species. We have not yet resolved whether wing size evolution at the <em>ws1</em> locus is caused by regulatory alterations of <em>dsx</em> or <em>prospero,</em> or by another mechanism. This study demonstrates the feasibility of efficient positional cloning of quantitative trait loci (QTL) involved in a broad array of phenotypic differences among <em>Nasonia</em> species.</p></div>", "links"=>[], "tags"=>["non-coding", "changes", "sex-specific", "differences"], "article_id"=>144995, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1000821.s001", "https://dx.doi.org/10.1371/journal.pgen.1000821.s002", "https://dx.doi.org/10.1371/journal.pgen.1000821.s003", "https://dx.doi.org/10.1371/journal.pgen.1000821.s004"], "stats"=>{"downloads"=>0, "page_views"=>22, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Non_Coding_Changes_Cause_Sex_Specific_Wing_Size_Differences_between_Closely_Related_Species_of_Nasonia_/144995", "title"=>"Non-Coding Changes Cause Sex-Specific Wing Size Differences between Closely Related Species of <em>Nasonia</em>", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2010-01-15 01:23:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/867644"], "description"=>"<p>Seta area and seta number are used to estimate cell size and cell number in the distal portion of the forewing. Seta area is the mean area occupied by each seta, based on the distance to each seta's nearest four neighbors. Absolute measurements (mean ± standard deviation) are shown for <i>ws1<sub>v</sub>V</i> (<i>N. vitripennis</i>) males (all area units in µm<sup>2</sup>). Relative measurements are shown as (mean / <i>ws1<sub>v</sub>V</i> mean) ± (standard deviation / <i>ws1<sub>v</sub>V</i> mean). N: number of individuals measured, nested in (families).</p>", "links"=>[], "tags"=>["seta"], "article_id"=>538093, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.t002", "stats"=>{"downloads"=>1, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Cell_number_and_size_effects_of_ws1_as_estimated_by_seta_number_and_area_/538093", "title"=>"Cell number and size effects of <i>ws1</i>, as estimated by seta number and area.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-01-15 02:14:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/867605"], "description"=>"<p>Absolute measurements (mean ± standard deviation) for <i>ws1<sub>v</sub>V</i> (<i>N. vitripennis</i>) males and females are shown (length and width in µm, area in µm<sup>2</sup>). Relative measurements for other genotypes are shown as (mean / <i>ws1<sub>v</sub>V</i> mean) ± (standard deviation / <i>ws1<sub>v</sub>V</i> mean). <i>a,b,c</i>: contrast groups for multiple comparisons (Tukey's HSD) between genotypes within each sex. Groups are unchanged for confidence levels of both alpha = 0.05 and 0.001. N: number of individuals measured, nested in (families).</p>", "links"=>[], "tags"=>["wild-type"], "article_id"=>538058, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.t001", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Basic_wing_measurements_of_ws1_and_wild_type_strains_/538058", "title"=>"Basic wing measurements of <i>ws1</i> and wild-type strains.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-01-15 02:14:18"}
  • {"files"=>["https://ndownloader.figshare.com/files/867202"], "description"=>"<p>Wings of <i>N. giraulti</i> (<i>ws1<sub>g</sub>G</i>), <i>N. vitripennis</i> (<i>ws1<sub>v</sub>V</i>) and <i>giraulti ws1</i> in <i>vitripennis</i> background (<i>ws1<sub>g</sub>V_40kb</i>). Wing area ± S.D. relative to <i>ws1<sub>v</sub>V</i> males is shown (see also <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1000821#pgen-1000821-t001\" target=\"_blank\">Table 1</a>). Scale bar: 100 µm.</p>", "links"=>[], "tags"=>["differences"], "article_id"=>537654, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.g001", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Wing_size_differences_due_to_ws1_/537654", "title"=>"Wing size differences due to <i>ws1.</i>", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-01-15 02:07:34"}
  • {"files"=>["https://ndownloader.figshare.com/files/867689"], "description"=>"<p>Expression level change was estimated by quantitative RT-PCR. Expression ratios greater than 1 indicate higher transcript level in <i>ws1<sub>v</sub>V</i> than in <i>ws1<sub>g</sub>V_40kb</i>, corrected for control gene (<i>rp49</i>) expression. N: number of independent biological replicates. <i>dsx<sup>M</sup></i>: male <i>dsx</i> splice form. <i>dsx</i> in females used non-sex-specific primers.</p>", "links"=>[], "tags"=>["developmental biology/developmental evolution", "evolutionary biology/developmental evolution", "evolutionary biology/evolutionary and comparative genetics", "evolutionary biology/morphogenesis and cell biology", "genetics and genomics/animal genetics", "genetics and genomics/complex traits", "genetics and genomics/gene discovery"], "article_id"=>538131, "categories"=>["Genetics", "Evolutionary Biology", "Developmental Biology"], "users"=>["David W. Loehlin", "Deodoro C. S. G. Oliveira", "Rachel Edwards", "Jonathan D. Giebel", "Michael E. Clark", "M. Victoria Cattani", "Louis van de Zande", "Eveline C. Verhulst", "Leo W. Beukeboom", "Monica Muñoz-Torres", "John H. Werren"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000821.t003", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Gene_expression_analysis_in_the_ws1_region_/538131", "title"=>"Gene expression analysis in the <i>ws1</i> region.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-01-15 02:15:31"}

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  • {"unique-ip"=>"18", "full-text"=>"16", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"4", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2014", "month"=>"11"}
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  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"7"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"8"}
  • {"unique-ip"=>"10", "full-text"=>"9", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2015", "month"=>"9"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"10"}
  • {"unique-ip"=>"7", "full-text"=>"5", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2015", "month"=>"11"}
  • {"unique-ip"=>"6", "full-text"=>"4", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"12"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"1"}
  • {"unique-ip"=>"9", "full-text"=>"6", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"2"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"3"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"4"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"5"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"6"}
  • {"unique-ip"=>"3", "full-text"=>"1", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"7"}
  • {"unique-ip"=>"1", "full-text"=>"2", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"8"}
  • {"unique-ip"=>"5", "full-text"=>"2", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"11"}
  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"12"}
  • {"unique-ip"=>"5", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2017", "month"=>"1"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"2"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"3"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"4"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"5"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"6"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"7"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"8"}
  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2017", "month"=>"9"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"11"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"12"}
  • {"unique-ip"=>"22", "full-text"=>"12", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"5", "cited-by"=>"1", "year"=>"2018", "month"=>"1"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"3"}
  • {"unique-ip"=>"7", "full-text"=>"7", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"1"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"5"}
  • {"unique-ip"=>"10", "full-text"=>"7", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"6"}
  • {"unique-ip"=>"4", "full-text"=>"1", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"7", "full-text"=>"9", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"7", "full-text"=>"8", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"4", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2018", "month"=>"11"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"2"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"4"}
  • {"unique-ip"=>"4", "full-text"=>"2", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}

Relative Metric

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