Fis Is Essential for Capsule Production in Pasteurella multocida and Regulates Expression of Other Important Virulence Factors
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{"title"=>"Fis is essential for capsule production in Pasteurella multocida and regulates expression of other important virulence factors", "type"=>"journal", "authors"=>[{"first_name"=>"Jason A.", "last_name"=>"Steen", "scopus_author_id"=>"51664080500"}, {"first_name"=>"Jennifer A.", "last_name"=>"Steen", "scopus_author_id"=>"26536706900"}, {"first_name"=>"Paul", "last_name"=>"Harrison", "scopus_author_id"=>"24173346400"}, {"first_name"=>"Torsten", "last_name"=>"Seemann", "scopus_author_id"=>"35273114400"}, {"first_name"=>"Ian", "last_name"=>"Wilkie", "scopus_author_id"=>"7004156705"}, {"first_name"=>"Marina", "last_name"=>"Harper", "scopus_author_id"=>"7201849193"}, {"first_name"=>"Ben", "last_name"=>"Adler", "scopus_author_id"=>"7102049681"}, {"first_name"=>"John D.", "last_name"=>"Boyce", "scopus_author_id"=>"7201747360"}], "year"=>2010, "source"=>"PLoS Pathogens", "identifiers"=>{"pmid"=>"20140235", "sgr"=>"77649265411", "scopus"=>"2-s2.0-77649265411", "isbn"=>"1553-7366", "doi"=>"10.1371/journal.ppat.1000750", "pui"=>"358395482", "issn"=>"15537366"}, "id"=>"c601f5da-ba85-306a-825e-e06d614dcfa3", "abstract"=>"P. multocida is the causative agent of a wide range of diseases of animals, including fowl cholera in poultry and wild birds. Fowl cholera isolates of P. multocida generally express a capsular polysaccharide composed of hyaluronic acid. There have been reports of spontaneous capsule loss in P. multocida, but the mechanism by which this occurs has not been determined. In this study, we identified three independent strains that had spontaneously lost the ability to produce capsular polysaccharide. Quantitative RT-PCR showed that these strains had significantly reduced transcription of the capsule biosynthetic genes, but DNA sequence analysis identified no mutations within the capsule biosynthetic locus. However, whole-genome sequencing of paired capsulated and acapsular strains identified a single point mutation within the fis gene in the acapsular strain. Sequencing of fis from two independently derived spontaneous acapsular strains showed that each contained a mutation within fis. Complementation of these strains with an intact copy of fis, predicted to encode a transcriptional regulator, returned capsule expression to all strains. Therefore, expression of a functional Fis protein is essential for capsule expression in P. multocida. DNA microarray analysis of one of the spontaneous fis mutants identified approximately 30 genes as down-regulated in the mutant, including pfhB_2, which encodes a filamentous hemagglutinin, a known P. multocida virulence factor, and plpE, which encodes the cross protective surface antigen PlpE. Therefore these experiments define for the first time a mechanism for spontaneous capsule loss in P. multocida and identify Fis as a critical regulator of capsule expression. Furthermore, Fis is involved in the regulation of a range of other P. multocida genes including important virulence factors.", "link"=>"http://www.mendeley.com/research/fis-essential-capsule-production-pasteurella-multocida-regulates-expression-other-important-virulenc", "reader_count"=>35, "reader_count_by_academic_status"=>{"Researcher"=>15, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>6, "Student > Master"=>5, "Other"=>1, "Student > Bachelor"=>4, "Lecturer"=>1}, "reader_count_by_user_role"=>{"Researcher"=>15, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>6, "Student > Master"=>5, "Other"=>1, "Student > Bachelor"=>4, "Lecturer"=>1}, "reader_count_by_subject_area"=>{"Biochemistry, Genetics and Molecular Biology"=>2, "Agricultural and Biological Sciences"=>27, "Medicine and Dentistry"=>3, "Design"=>1, "Veterinary Science and Veterinary Medicine"=>1, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Design"=>{"Design"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>3}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>27}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>2}, "Veterinary Science and Veterinary Medicine"=>{"Veterinary Science and Veterinary Medicine"=>1}}, "reader_count_by_country"=>{"Argentina"=>1, "United States"=>4, "Denmark"=>1, "Brazil"=>2}, "group_count"=>3}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/863790"], "description"=>"<p>The amount of capsular material is shown for paired acapsular (AL1114, AL1162 and AL1396) and capsulated (AL1115, AL1163 and AL1397) variants and the same strains complemented with either empty vector (AL1398, AL1400 and AL1402) or an intact copy of <i>fis</i> (AL1399, AL1401 and AL1403). Brackets below the figure indicate strains derived from the same parent strain. Also shown is the amount of HA produced by the TargeTron <i>fis</i> mutants AL1404 and AL1405, the wild-type encapsulated parent strain VP161 and the <i>hyaB</i> capsule mutant AL919. Results for experimentally related strains are shaded similarly with black bars representing <i>fis</i> mutants, white bars representing parent and wild-type strains, hatched bars representing the <i>fis</i> complemented strains, gray bars representing strains transformed with vector only and the dotted bar representing the directed <i>hyaB</i> mutant. Each bar represents triplicate values ±1 standard error of the mean (SEM) *** p&lt;0.001, ** p&lt;0.01, * p&lt;0.05.</p>", "links"=>[], "tags"=>["capsular", "polysaccharide", "produced"], "article_id"=>534238, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Hyaluronic_acid_HA_capsular_polysaccharide_produced_by_i_P_multocida_i_strains_/534238", "title"=>"Hyaluronic acid (HA) capsular polysaccharide produced by <i>P. multocida</i> strains.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 02:43:49"}
  • {"files"=>["https://ndownloader.figshare.com/files/863918"], "description"=>"<p>The organization of the genes in the capsule biosynthetic locus is shown in panel A and the nucleotide sequence of the intergenic region between <i>phyA</i> and <i>hyaE</i> is shown in the flyout box. The positions of the HyaE and PhyA translational initiation codons (ATG) are designated by right-angled arrowheads and the identified transcriptional start site for <i>hyaE</i> is shown in bold text and labeled +1. Predicted RBS, −10 and −35 sites are underlined, and labeled above the sequence. Predicted Fis binding sites based on the <i>E. coli</i> position specific weight matrix are boxed <a href=\"http://www.plospathogens.org/article/info:doi/10.1371/journal.ppat.1000750#ppat.1000750-Bradley1\" target=\"_blank\">[15]</a>. The length (x-axis) and quantity (y-axis) of <i>hyaE</i>-specific primer extension products generated using VP161 total RNA and the primer BAP5476 are shown in panel B. The exact size (in bp) of the major primer extension products are shown beside the peaks.</p>", "links"=>[], "tags"=>["serogroup", "capsule", "biosynthetic", "locus", "transcriptional"], "article_id"=>534362, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Schematic_representation_of_the_i_P_multocida_i_serogroup_A_capsule_biosynthetic_locus_and_identification_of_the_i_hyaE_i_transcriptional_start_site_/534362", "title"=>"Schematic representation of the <i>P. multocida</i> serogroup A capsule biosynthetic locus and identification of the <i>hyaE</i> transcriptional start site.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 02:44:35"}
  • {"files"=>["https://ndownloader.figshare.com/files/864021"], "description"=>"<p>The expression levels of <i>phyA</i> (panel A) and <i>hyaE</i> (panel B) were determined for the strains VP161 (wild-type), AL1114 (spontaneous acapsular strain), AL1115 (paired capsulated strain) and AL919 (a VP161 <i>hyaB</i> insertional mutant) by qRT-PCR and normalized to the level of <i>gyrB</i> expression. The average relative expression was determined from three biological replicates and the values shown are the mean ±1 SEM. ** p&lt;0.01, * p&lt;0.05.</p>", "links"=>[], "tags"=>["biochemistry/bioinformatics", "microbiology/cellular microbiology and pathogenesis", "microbiology/immunity to infections"], "article_id"=>534473, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Relative_expression_of_i_phyA_i_and_i_hyaE_i_in_four_i_P_multocida_i_strains_/534473", "title"=>"Relative expression of <i>phyA</i> and <i>hyaE</i> in four <i>P. multocida</i> strains.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 02:45:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/864107"], "description"=>"<p>The putative promoter-containing regions upstream of <i>hyaE</i>, <i>phyA</i>, <i>pglA</i>, <i>pm0998</i>, <i>pm1078</i>, <i>lspB_2</i>, and <i>pm1818</i> were cloned into the <i>P. multocida</i> promoter detecting vector pMKΩ, and the ability of the recombinant constructs (pAL597, pAL596, pAL795, pAL796, pAL797, pAL798 and pAL799 respectively) to confer kanamycin resistance to strains AL1114 (spontaneous acapsular strain), AL1115 (paired capsulated strain) and VP161 (capsulated wild-type parent) assessed. Empty pMKΩ vector was used as a negative control. The average relative expression was determined from three biological replicates and the values shown are the mean ±1 SEM. *** p&lt;0.001, ** p&lt;0.01.</p>", "links"=>[], "tags"=>["strains", "kanamycin", "diffusion"], "article_id"=>534559, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Sensitivity_of_i_P_multocida_i_strains_to_kanamycin_as_determined_by_disc_diffusion_assays_/534559", "title"=>"Sensitivity of <i>P. multocida</i> strains to kanamycin as determined by disc diffusion assays.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 02:45:42"}
  • {"files"=>["https://ndownloader.figshare.com/files/864203"], "description"=>"<p>Western immunoblot using chicken antiserum against recombinant PlpE (A) and the corresponding quantitive densitometry analysis (B). The levels of PlpE expression were measured in the capsulated parent strains (VP161, AL435), the acapsular <i>fis</i> mutants (AL1114, AL1162) and paired capsulated strains (AL1115, AL1163), a single-crossover <i>plpE</i> mutant (AL1172; Hatfaludi, Al-Hasani and Adler, unpublished data), complemented <i>plpE</i> mutant (AL1174) and the acapsular <i>phyB</i> mutant strain that produces no HA (AL1164). The filled arrow head represents the position of PlpE, whilst the unfilled arrowhead represents the background band used as a loading control for densitometry normalization. Each bar represents the average normalized densitometry readings from triplicate biological replicates ±1 SEM. ** p&lt;0.01, * p&lt;0.05.</p>", "links"=>[], "tags"=>["immunoblot", "densitometry", "plpe", "capsulated"], "article_id"=>534652, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Western_immunoblot_and_densitometry_analysis_of_i_P_multocida_i_PlpE_expression_in_acapsular_capsulated_and_control_strains_/534652", "title"=>"Western immunoblot and densitometry analysis of <i>P. multocida</i> PlpE expression in acapsular, capsulated and control strains.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 02:46:14"}
  • {"files"=>["https://ndownloader.figshare.com/files/864277"], "description"=>"a<p>restriction sites are underlined.</p>", "links"=>[], "tags"=>["biochemistry/bioinformatics", "microbiology/cellular microbiology and pathogenesis", "microbiology/immunity to infections"], "article_id"=>534730, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.t002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Oligonucleotides_used_in_this_study_/534730", "title"=>"Oligonucleotides used in this study.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 02:46:40"}
  • {"files"=>["https://ndownloader.figshare.com/files/864322"], "description"=>"a<p>All sequencing reads were aligned to the annotated <i>P. multocida</i> Pm70 genome sequence and the position of each mutation in relation to this sequence shown.</p>", "links"=>[], "tags"=>["differences", "whole-genome", "illumina", "sequencing", "acapsular", "al1114", "paired", "capsulated"], "article_id"=>534766, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.t003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Sequence_differences_identified_by_whole_genome_Illumina_short_read_sequencing_between_the_acapsular_strain_AL1114_and_the_paired_capsulated_strain_AL1115_/534766", "title"=>"Sequence differences identified by whole-genome Illumina short read sequencing between the acapsular strain AL1114 and the paired capsulated strain AL1115.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 02:46:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/864363"], "description"=>"<p>Strains and plasmids used in this study.</p>", "links"=>[], "tags"=>["plasmids"], "article_id"=>534820, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.t001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Strains_and_plasmids_used_in_this_study_/534820", "title"=>"Strains and plasmids used in this study.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 02:47:08"}
  • {"files"=>["https://ndownloader.figshare.com/files/864401"], "description"=>"a<p>NCBI accession number of the identified gene.</p>b<p>Cluster of Orthologous Groups functional group (<a href=\"http://www.ncbi.nlm.nih.gov/COG/\" target=\"_blank\">http://www.ncbi.nlm.nih.gov/COG/</a>).</p>c<p>Only one of two probes for this gene was significantly differentially regulated.</p>", "links"=>[], "tags"=>["differentially", "regulated", "wild-type", "mutant", "dna", "microarray"], "article_id"=>534854, "categories"=>["Biochemistry", "Microbiology"], "users"=>["Jason A. Steen", "Jennifer A. Steen", "Paul Harrison", "Torsten Seemann", "Ian Wilkie", "Marina Harper", "Ben Adler", "John D. Boyce"], "doi"=>["https://dx.doi.org/10.1371/journal.ppat.1000750.t004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Genes_identified_as_differentially_regulated_between_the_i_P_multocida_i_wild_type_strain_and_a_i_fis_i_mutant_strain_as_determined_by_DNA_microarray_analysis_/534854", "title"=>"Genes identified as differentially regulated between the <i>P. multocida</i> wild-type strain and a <i>fis</i> mutant strain as determined by DNA microarray analysis.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-21 02:47:19"}

PMC Usage Stats | Further Information

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  • {"unique-ip"=>"11", "full-text"=>"9", "pdf"=>"5", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"12"}
  • {"unique-ip"=>"13", "full-text"=>"14", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"1"}
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  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"3"}
  • {"unique-ip"=>"12", "full-text"=>"13", "pdf"=>"7", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"4"}
  • {"unique-ip"=>"12", "full-text"=>"25", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"16", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"5"}
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  • {"unique-ip"=>"7", "full-text"=>"6", "pdf"=>"9", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"11", "full-text"=>"10", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
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  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"12"}
  • {"unique-ip"=>"7", "full-text"=>"6", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"1"}
  • {"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"=>"2"}
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  • {"unique-ip"=>"7", "full-text"=>"9", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"9"}
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  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"4", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"11"}
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  • {"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"=>"2018", "month"=>"1"}
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  • {"unique-ip"=>"7", "full-text"=>"9", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
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  • {"unique-ip"=>"5", "full-text"=>"3", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
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  • {"unique-ip"=>"10", "full-text"=>"9", "pdf"=>"8", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"12", "full-text"=>"14", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"8", "full-text"=>"7", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"11"}
  • {"unique-ip"=>"11", "full-text"=>"5", "pdf"=>"5", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"9", "full-text"=>"10", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"2"}
  • {"unique-ip"=>"6", "full-text"=>"7", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"4"}
  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}
  • {"unique-ip"=>"30", "full-text"=>"39", "pdf"=>"5", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"14", "full-text"=>"15", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
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Relative Metric

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