G-Protein Coupled Receptor Signaling Architecture of Mammalian Immune Cells
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{"title"=>"G-protein coupled receptor signaling architecture of mammalian immune cells", "type"=>"journal", "authors"=>[{"first_name"=>"Natalia", "last_name"=>"Polouliakh", "scopus_author_id"=>"8925116000"}, {"first_name"=>"Richard", "last_name"=>"Nock", "scopus_author_id"=>"6701497179"}, {"first_name"=>"Frank", "last_name"=>"Nielsen", "scopus_author_id"=>"55167047900"}, {"first_name"=>"Hiroaki", "last_name"=>"Kitano", "scopus_author_id"=>"7201618005"}], "year"=>2009, "source"=>"PLoS ONE", "identifiers"=>{"doi"=>"10.1371/journal.pone.0004189", "sgr"=>"58449099653", "issn"=>"19326203", "pui"=>"354084421", "isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)", "pmid"=>"19142232", "scopus"=>"2-s2.0-58449099653"}, "id"=>"35972ccc-b6b4-3ee6-ab7f-4c4d480bf438", "abstract"=>"A series of recent studies on large-scale networks of signaling and metabolic systems revealed that a certain network structure often called \"bow-tie network\" are observed. In signaling systems, bow-tie network takes a form with diverse and redundant inputs and outputs connected via a small numbers of core molecules. While arguments have been made that such network architecture enhances robustness and evolvability of biological systems, its functional role at a cellular level remains obscure. A hypothesis was proposed that such a network function as a stimuli-reaction classifier where dynamics of core molecules dictate downstream transcriptional activities, hence physiological responses against stimuli. In this study, we examined whether such hypothesis can be verified using experimental data from Alliance for Cellular Signaling (AfCS) that comprehensively measured GPCR related ligands response for B-cell and macrophage. In a GPCR signaling system, cAMP and Ca2+ act as core molecules. Stimuli-response for 32 ligands to B-Cells and 23 ligands to macrophages has been measured. We found that ligands with correlated changes of cAMP and Ca2+ tend to cluster closely together within the hyperspaces of both cell types and they induced genes involved in the same cellular processes. It was found that ligands inducing cAMP synthesis activate genes involved in cell growth and proliferation; cAMP and Ca2+ molecules that increased together form a feedback loop and induce immune cells to migrate and adhere together. In contrast, ligands without a core molecules response are scattered throughout the hyperspace and do not share clusters. G-protein coupling receptors together with immune response specific receptors were found in cAMP and Ca2+ activated clusters. Analyses have been done on the original software applicable for discovering 'bow-tie' network architectures within the complex network of intracellular signaling where ab initio clustering has been implemented as well. Groups of potential transcription factors for each specific group of genes were found to be partly conserved across B-cell and macrophage. A series of findings support the hypothesis.", "link"=>"http://www.mendeley.com/research/gprotein-coupled-receptor-signaling-architecture-mammalian-immune-cells-1", "reader_count"=>43, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>13, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>2, "Student > Master"=>1, "Other"=>2, "Lecturer"=>1, "Professor"=>2}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>13, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>2, "Student > Master"=>1, "Other"=>2, "Lecturer"=>1, "Professor"=>2}, "reader_count_by_subject_area"=>{"Engineering"=>4, "Unspecified"=>2, "Biochemistry, Genetics and Molecular Biology"=>1, "Materials Science"=>1, "Agricultural and Biological Sciences"=>30, "Medicine and Dentistry"=>1, "Chemistry"=>1, "Computer Science"=>2, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>4}, "Materials Science"=>{"Materials Science"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Chemistry"=>{"Chemistry"=>1}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>30}, "Computer Science"=>{"Computer Science"=>2}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>1}, "Unspecified"=>{"Unspecified"=>2}}, "reader_count_by_country"=>{"United States"=>2, "Japan"=>1, "United Kingdom"=>3}, "group_count"=>1}

Scopus | Further Information

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  • {"files"=>["https://ndownloader.figshare.com/files/911371"], "description"=>"<p>Two cAMP-activated ligands (PGE and ISO) form a distinct <i>sub</i>-region within the hyperspace. NO/NO ligands (LPS and IFG) do not form a single <i>sub</i>-region. 2MA is an only YES/YES ligand remoted from YES/NO <i>sub</i>-region as well as from NO/NO ligands. Possibly Cluster_8 is the influenced by 2MA. This representation, which we call ‘PCA correlation ball’, shows both the groups of ligands and their correlations, and also the regulations of gene clusters from the projections of nearby ligands. Genes functions (‘Function’ or ‘Cellular process’ (GO terms), if ‘Function’ is absent) calculated basing on the enrichment in clusters with p-value<0.05 are shown in the pie-graphs. By the reason of small number of genes we combined <i>up-</i> and <i>down-</i> regulated gene together in the pie-graphs.</p>", "links"=>[], "tags"=>["macrophage", "ligands", "microarray", "iso", "2ma", "ifg"], "article_id"=>581824, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.g005", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Clustering_results_on_5_macrophage_ligands_with_available_microarray_expression_data_PGE_ISO_YES_NO_2MA_YES_YES_and_LPS_IFG_NO_NO_/581824", "title"=>"Clustering results on 5 macrophage ligands with available microarray expression data: PGE, ISO (YES/NO), 2MA (YES/YES) and LPS, IFG (NO/NO).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-01-14 00:30:24"}
  • {"files"=>["https://ndownloader.figshare.com/files/911497"], "description"=>"<p>Receptor genes expressed in YES/NO and NO/YES <i>sub</i>-regions of macrophage.</p>", "links"=>[], "tags"=>["genes"], "article_id"=>581952, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.t002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Receptor_genes_expressed_in_YES_NO_and_NO_YES_sub_regions_of_macrophage_/581952", "title"=>"Receptor genes expressed in YES/NO and NO/YES <i>sub</i>-regions of macrophage.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-01-14 00:32:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/911190"], "description"=>"<p>Standard deviations (SD) are shown by the vertical bars. Full names for the ligands are listed in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004189#pone.0004189.s004\" target=\"_blank\">Figure S2</a>. Values equal to zero for cAMP correspond to the experimentally uncharacterized cAMP. 2MA ligand is marked to be visible as it is given more detailed discussion in the manuscript.</p>", "links"=>[], "tags"=>["changes", "23", "macrophage", "ligands", "classified", "groups", "observed", "non-observed", "elevation"], "article_id"=>581636, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.g003", "stats"=>{"downloads"=>1, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Peaks_of_fold_changes_for_23_macrophage_ligands_classified_into_three_groups_with_observed_YES_and_non_observed_NO_elevation_of_Ca_2_and_cAMP_/581636", "title"=>"Peaks of fold changes for 23 macrophage ligands classified into three groups with observed ‘YES’ and non-observed ‘NO’ elevation of Ca<sup>2+</sup> and cAMP.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-01-14 00:27:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/911096"], "description"=>"<p>Standard deviations (SD) are shown by the vertical bars. Full names for the ligands are listed in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004189#pone.0004189.s001\" target=\"_blank\">Table S1</a>. S1P and LPA ligands are marked to be visible as they are given more detailed discussion in the manuscript.</p>", "links"=>[], "tags"=>["changes", "32", "B-cell", "ligands", "classified", "groups", "observed", "non-observed", "elevation"], "article_id"=>581545, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.g002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Peaks_of_fold_changes_for_32_B_Cell_ligands_classified_into_four_groups_with_observed_YES_and_non_observed_NO_elevation_of_cAMP_and_Ca_2_/581545", "title"=>"Peaks of fold changes for 32 B-Cell ligands classified into four groups with observed ‘YES’ and non-observed ‘NO’ elevation of cAMP and Ca<sup>2+</sup>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-01-14 00:25:45"}
  • {"files"=>["https://ndownloader.figshare.com/files/911470"], "description"=>"<p>Receptor genes expressed in YES/NO and YES/YES <i>sub</i>-regions of B-Cell.</p>", "links"=>[], "tags"=>["genes"], "article_id"=>581930, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.t001", "stats"=>{"downloads"=>2, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Receptor_genes_expressed_in_YES_NO_and_YES_YES_sub_regions_of_B_Cell_/581930", "title"=>"Receptor genes expressed in YES/NO and YES/YES <i>sub</i>-regions of B-Cell.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-01-14 00:32:10"}
  • {"files"=>["https://ndownloader.figshare.com/files/450548", "https://ndownloader.figshare.com/files/450627", "https://ndownloader.figshare.com/files/450687", "https://ndownloader.figshare.com/files/450744", "https://ndownloader.figshare.com/files/450793", "https://ndownloader.figshare.com/files/450837"], "description"=>"<div><p>A series of recent studies on large-scale networks of signaling and metabolic systems revealed that a certain network structure often called “bow-tie network” are observed. In signaling systems, bow-tie network takes a form with diverse and redundant inputs and outputs connected via a small numbers of core molecules. While arguments have been made that such network architecture enhances robustness and evolvability of biological systems, its functional role at a cellular level remains obscure. A hypothesis was proposed that such a network function as a stimuli-reaction classifier where dynamics of core molecules dictate downstream transcriptional activities, hence physiological responses against stimuli. In this study, we examined whether such hypothesis can be verified using experimental data from Alliance for Cellular Signaling (AfCS) that comprehensively measured GPCR related ligands response for B-cell and macrophage. In a GPCR signaling system, cAMP and Ca<sup>2+</sup> act as core molecules. Stimuli-response for 32 ligands to B-Cells and 23 ligands to macrophages has been measured. We found that ligands with correlated changes of cAMP and Ca<sup>2+</sup> tend to cluster closely together within the hyperspaces of both cell types and they induced genes involved in the same cellular processes. It was found that ligands inducing cAMP synthesis activate genes involved in cell growth and proliferation; cAMP and Ca<sup>2+</sup> molecules that increased together form a feedback loop and induce immune cells to migrate and adhere together. In contrast, ligands without a core molecules response are scattered throughout the hyperspace and do not share clusters. G-protein coupling receptors together with immune response specific receptors were found in cAMP and Ca<sup>2+</sup> activated clusters. Analyses have been done on the original software applicable for discovering ‘bow-tie’ network architectures within the complex network of intracellular signaling where <em>ab initio</em> clustering has been implemented as well. Groups of potential transcription factors for each specific group of genes were found to be partly conserved across B-Cell and macrophage. A series of findings support the hypothesis.</p></div>", "links"=>[], "tags"=>["g-protein", "coupled", "receptor", "signaling", "mammalian", "cells"], "article_id"=>148809, "categories"=>["Cell Biology", "Biological Sciences"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0004189.s001", "https://dx.doi.org/10.1371/journal.pone.0004189.s002", "https://dx.doi.org/10.1371/journal.pone.0004189.s003", "https://dx.doi.org/10.1371/journal.pone.0004189.s004", "https://dx.doi.org/10.1371/journal.pone.0004189.s005", "https://dx.doi.org/10.1371/journal.pone.0004189.s006"], "stats"=>{"downloads"=>21, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/G_Protein_Coupled_Receptor_Signaling_Architecture_of_Mammalian_Immune_Cells/148809", "title"=>"G-Protein Coupled Receptor Signaling Architecture of Mammalian Immune Cells", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2009-01-14 02:26:49"}
  • {"files"=>["https://ndownloader.figshare.com/files/911015"], "description"=>"<p>Bow-tie networks generally have diverse inputs and outputs with conserved core nodes. It is called “bow-tie” as its pictorial representation resembles bowtie. Fig 1(a): In a three layer feed forward neural networks, a hidden layer (a middle layer) provides generalization capability and the neural network can function as classifier of diverse inputs mapping into diverse output patterns. It is essential that numbers of nodes in the hidden layer is smaller than numbers of nodes in inputs and output layers in order to achieve high level of generalization. This is an example of 6∶3∶6 feed forward network with 6 input nodes, 3 hidden nodes, and 6 output layer nodes. Fig 1(b): In signaling networks, receptors corresponds nodes in an input layer, molecules such as cAMP and calcium corresponds nodes in a hidden layer, and transcription factors corresponds to nodes in an output layer (Left). Looking at this from a stimuli-response viewpoint, it shall be viewed as a process that diverse ligands activate variety of receptors forming distinct activation patterns and results in patterns of activations at transcriptional level (Right). An intermediate layer (core nodes) that corresponds to the hidden layer in the feed forward neural network shall provide generalization capability to the signaling network. In GPCR pathway, cAMP and calcium are key molecules constituting this layer. Thus, diverse stimuli are classified into several groups that have similar calcium and cAMP elevation. Ligands that are classified into the same group shall activate similar subset of genes, hence invoking similar physiological responses, if generalization is actually taking place.</p>", "links"=>[], "tags"=>["cell biology/cell signaling", "cell biology/gene expression", "computational biology/transcriptional regulation"], "article_id"=>581469, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.g001", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Bow_tie_network_for_signaling_/581469", "title"=>"Bow-tie network for signaling.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-01-14 00:24:29"}
  • {"files"=>["https://ndownloader.figshare.com/files/911269"], "description"=>"<p>Ligands group into two big <i>sub</i>-regions: YES/NO and YES/YES and one small <i>sub</i>-region represented by MI3a (NO/YES) ligand. The M3A ligand mainly induced by Ca<sup>2+</sup> is separated and does not form a cluster downstream. This representation, which we call ‘PCA correlation ball’, shows both the groups of ligands and their correlations, and also the regulations of gene clusters from the projections of nearby ligands. Bow-graphs in the upper corners show the percent of genes being <i>up-</i> or <i>down-</i> regulated after the expiration of one/both molecules activation limited to first time marks (30 min in B-Cell), and the differences are obvious. Genes functions (‘Function’ or ‘Cellular process’ (GO terms), if ‘Function’ is absent) calculated basing on the enrichment in clusters with p-value<0.05 are shown in the pie-graphs.</p>", "links"=>[], "tags"=>["cell biology/cell signaling", "cell biology/gene expression", "computational biology/transcriptional regulation"], "article_id"=>581726, "categories"=>["Cell Biology", "Computational Biology"], "users"=>["Natalia Polouliakh", "Richard Nock", "Frank Nielsen", "Hiroaki Kitano"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0004189.g004", "stats"=>{"downloads"=>1, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Clustering_results_for_YES_NO_cAMP_YES_YES_cAMP_Ca_2_and_NO_YES_Ca_2_groups_/581726", "title"=>"Clustering results for YES/NO (cAMP), YES/YES (cAMP-Ca<sup>2+</sup>) and NO/YES (Ca<sup>2+</sup>) groups.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-01-14 00:28:46"}

PMC Usage Stats | Further Information

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  • {"unique-ip"=>"12", "full-text"=>"7", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"7", "supp-data"=>"4", "cited-by"=>"2", "year"=>"2014", "month"=>"9"}
  • {"unique-ip"=>"16", "full-text"=>"12", "pdf"=>"6", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"7", "supp-data"=>"6", "cited-by"=>"1", "year"=>"2014", "month"=>"10"}
  • {"unique-ip"=>"2", "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"=>"2"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2014", "month"=>"11"}
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  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"1"}
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  • {"unique-ip"=>"2", "full-text"=>"1", "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"=>"1", "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"=>"4"}
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  • {"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"=>"2016", "month"=>"6"}
  • {"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"=>"7"}
  • {"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"=>"8"}
  • {"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"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"2", "full-text"=>"3", "pdf"=>"0", "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"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"12"}
  • {"unique-ip"=>"6", "full-text"=>"4", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"1"}
  • {"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"=>"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"=>"2", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"4"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"5"}
  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"3", "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"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"7"}
  • {"unique-ip"=>"6", "full-text"=>"4", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"8"}
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  • {"unique-ip"=>"7", "full-text"=>"6", "pdf"=>"4", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
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  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"12"}
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  • {"unique-ip"=>"6", "full-text"=>"6", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"3"}
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  • {"unique-ip"=>"8", "full-text"=>"4", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"5", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"14", "full-text"=>"12", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"11", "full-text"=>"11", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"6", "full-text"=>"5", "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"=>"4", "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"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
  • {"unique-ip"=>"1", "full-text"=>"0", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"4"}
  • {"unique-ip"=>"6", "full-text"=>"6", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}
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Relative Metric

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