A Minimal Model of Metabolism-Based Chemotaxis
Publication Date
December 02, 2010
Journal
PLOS Computational Biology
Authors
Matthew D. Egbert, Xabier E. Barandiaran & Ezequiel A. Di Paolo
Volume
6
Issue
12
Pages
e1001004
DOI
https://dx.plos.org/10.1371/journal.pcbi.1001004
Publisher URL
http://journals.plos.org/ploscompbiol/article?id=10.1371%2Fjournal.pcbi.1001004
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/21170312
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3000427
Europe PMC
http://europepmc.org/abstract/MED/21170312
Web of Science
000285574600002
Scopus
78651244766
Mendeley
http://www.mendeley.com/research/minimal-model-metabolismbased-chemotaxis
Events
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Mendeley | Further Information

{"title"=>"A minimal model of metabolism-based chemotaxis", "type"=>"journal", "authors"=>[{"first_name"=>"Matthew D.", "last_name"=>"Egbert", "scopus_author_id"=>"35096042700"}, {"first_name"=>"Xabier E.", "last_name"=>"Barandiaran", "scopus_author_id"=>"6507085396"}, {"first_name"=>"Ezequiel A.", "last_name"=>"di Paolo", "scopus_author_id"=>"8878196500"}], "year"=>2010, "source"=>"PLoS Computational Biology", "identifiers"=>{"issn"=>"1553734X", "scopus"=>"2-s2.0-78651244766", "sgr"=>"78651244766", "pui"=>"361077264", "isbn"=>"1553-7358 (Electronic)\\r1553-734X (Linking)", "pmid"=>"21170312", "doi"=>"10.1371/journal.pcbi.1001004"}, "id"=>"3cd9dbd5-1217-38b8-aee2-9d972e71a4da", "abstract"=>"Since the pioneering work by Julius Adler in the 1960's, bacterial chemotaxis has been predominantly studied as metabolism-independent. All available simulation models of bacterial chemotaxis endorse this assumption. Recent studies have shown, however, that many metabolism-dependent chemotactic patterns occur in bacteria. We hereby present the simplest artificial protocell model capable of performing metabolism-based chemotaxis. The model serves as a proof of concept to show how even the simplest metabolism can sustain chemotactic patterns of varying sophistication. It also reproduces a set of phenomena that have recently attracted attention on bacterial chemotaxis and provides insights about alternative mechanisms that could instantiate them. We conclude that relaxing the metabolism-independent assumption provides important theoretical advances, forces us to rethink some established pre-conceptions and may help us better understand unexplored and poorly understood aspects of bacterial chemotaxis.", "link"=>"http://www.mendeley.com/research/minimal-model-metabolismbased-chemotaxis", "reader_count"=>71, "reader_count_by_academic_status"=>{"Unspecified"=>1, "Professor > Associate Professor"=>5, "Student > Doctoral Student"=>1, "Researcher"=>17, "Student > Ph. D. Student"=>27, "Student > Postgraduate"=>6, "Student > Master"=>6, "Student > Bachelor"=>1, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_user_role"=>{"Unspecified"=>1, "Professor > Associate Professor"=>5, "Student > Doctoral Student"=>1, "Researcher"=>17, "Student > Ph. D. Student"=>27, "Student > Postgraduate"=>6, "Student > Master"=>6, "Student > Bachelor"=>1, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_subject_area"=>{"Unspecified"=>3, "Agricultural and Biological Sciences"=>29, "Philosophy"=>9, "Computer Science"=>4, "Engineering"=>5, "Environmental Science"=>1, "Biochemistry, Genetics and Molecular Biology"=>8, "Mathematics"=>1, "Physics and Astronomy"=>5, "Psychology"=>2, "Social Sciences"=>2, "Immunology and Microbiology"=>1, "Linguistics"=>1}, "reader_count_by_subdiscipline"=>{"Social Sciences"=>{"Social Sciences"=>2}, "Physics and Astronomy"=>{"Physics and Astronomy"=>5}, "Psychology"=>{"Psychology"=>2}, "Mathematics"=>{"Mathematics"=>1}, "Unspecified"=>{"Unspecified"=>3}, "Environmental Science"=>{"Environmental Science"=>1}, "Engineering"=>{"Engineering"=>5}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>29}, "Computer Science"=>{"Computer Science"=>4}, "Linguistics"=>{"Linguistics"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>8}, "Philosophy"=>{"Philosophy"=>9}}, "reader_count_by_country"=>{"Netherlands"=>1, "United States"=>3, "Japan"=>1, "United Kingdom"=>4, "Germany"=>3, "India"=>1, "Spain"=>1, "Russia"=>1}, "group_count"=>5}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/814877"], "description"=>"<p>In a uniform distribution of resources, simulated bacteria move away from high concentrations of the metabolic inhibitor .</p>", "links"=>[], "tags"=>["metabolic", "inhibitors"], "article_id"=>485249, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g011", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_4_Metabolic_inhibitors_act_as_repellents_/485249", "title"=>"Experiment 4: Metabolic inhibitors act as repellents.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:02:50"}
  • {"files"=>["https://ndownloader.figshare.com/files/815078"], "description"=>"<p>The metabolism of the bacteria changes when encountering (located at the left side of the environment), as is incorporated into the cell, making metabolizable. The rich area (concentric circles on right side of plots) is initially insufficient to support production of , but for agents that have incorporated into their metabolism, becomes an attractant. Bacteria with are shown as s rather than circles.</p>", "links"=>[], "tags"=>["cell biology/chemical biology of the cell", "computational biology/metabolic networks", "computational biology/systems biology", "microbiology/microbial physiology and metabolism"], "article_id"=>485444, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g013", "stats"=>{"downloads"=>1, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_5_Sensitivity_to_history_/485444", "title"=>"Experiment 5: Sensitivity to history.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:03:50"}
  • {"files"=>["https://ndownloader.figshare.com/files/814436"], "description"=>"<p>Diagram indicating the minimal metabolic reaction and its coupling to tumbling and running modes of behavior.</p>", "links"=>[], "tags"=>["metabolic", "coupling", "tumbling", "modes"], "article_id"=>484801, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g005", "stats"=>{"downloads"=>2, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Diagram_indicating_the_minimal_metabolic_reaction_and_its_coupling_to_tumbling_and_running_modes_of_behavior_/484801", "title"=>"Diagram indicating the minimal metabolic reaction and its coupling to tumbling and running modes of behavior.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:00:28"}
  • {"files"=>["https://ndownloader.figshare.com/files/814499"], "description"=>"<p>Plot A is a histogram that indicates the distance of bacteria from the location of highest concentration of and () at the start and end of trials. Data are averaged from 10 runs of 100 bacteria each. Plot B indicates the spatial distribution of the simulated bacteria as time progresses in a typical trial. The concentric circles indicate the center of the Gaussian distribution of resources and .</p>", "links"=>[], "tags"=>["chemotaxis", "gradient"], "article_id"=>484873, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g006", "stats"=>{"downloads"=>1, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_1_Chemotaxis_towards_a_gradient_of_and_/484873", "title"=>"Experiment 1: Chemotaxis towards a gradient of and .", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:00:50"}
  • {"files"=>["https://ndownloader.figshare.com/files/814380"], "description"=>"<p>Energy required for a reaction to take place.</p>", "links"=>[], "tags"=>["cell biology/chemical biology of the cell", "computational biology/metabolic networks", "computational biology/systems biology", "microbiology/microbial physiology and metabolism"], "article_id"=>484747, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g004", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Energy_required_for_a_reaction_to_take_place_/484747", "title"=>"Energy required for a reaction to take place.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:00:12"}
  • {"files"=>["https://ndownloader.figshare.com/files/815276"], "description"=>"<p>Constant parameters used in the model. Reaction and degradation rate-constants are determined according to formulas applied to the free-energies and activation energies which are hand-designed to adhere to the constraints inherent in our definition of a minimal metabolism and to display the phenomena of interest.</p>", "links"=>[], "tags"=>["cell biology/chemical biology of the cell", "computational biology/metabolic networks", "computational biology/systems biology", "microbiology/microbial physiology and metabolism"], "article_id"=>485644, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.t001", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Constants_/485644", "title"=>"Constants.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-20 22:04:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/814828"], "description"=>"<p>Diagram of the metabolic reactions for Experiment 3.</p>", "links"=>[], "tags"=>["metabolic", "reactions"], "article_id"=>485198, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g010", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Diagram_of_the_metabolic_reactions_for_Experiment_3_/485198", "title"=>"Diagram of the metabolic reactions for Experiment 3.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:02:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/814584"], "description"=>"<p>Diagram indicating metabolic reactions for Experiment 2.</p>", "links"=>[], "tags"=>["metabolic", "reactions"], "article_id"=>484951, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g007", "stats"=>{"downloads"=>1, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Diagram_indicating_metabolic_reactions_for_Experiment_2_/484951", "title"=>"Diagram indicating metabolic reactions for Experiment 2.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:01:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/814331"], "description"=>"<p>An exergonic reaction () is coupled to an endergonic material transforming reaction () such that system components are produced () that catalyze those reactions.</p>", "links"=>[], "tags"=>["cell biology/chemical biology of the cell", "computational biology/metabolic networks", "computational biology/systems biology", "microbiology/microbial physiology and metabolism"], "article_id"=>484698, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g003", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Conceptualization_of_a_minimal_metabolism_/484698", "title"=>"Conceptualization of a minimal metabolism.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 21:59:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/814651"], "description"=>"<p>In this trial, agents were placed on the same gradients as in <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1001004#pcbi-1001004-g006\" target=\"_blank\">Figure 6</a>, but with an additional uniform distribution of alternative metabolizable sources . The presence of these resources clearly inhibits chemotaxis to and .</p>", "links"=>[], "tags"=>["abundance", "resources", "inhibits", "chemotaxis"], "article_id"=>485022, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g008", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_2_Abundance_of_sufficient_resources_inhibits_chemotaxis_to_attractants_/485022", "title"=>"Experiment 2: Abundance of sufficient resources inhibits chemotaxis to attractants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:01:37"}
  • {"files"=>["https://ndownloader.figshare.com/files/815187"], "description"=>"<p>Here we see simulated bacteria concentrated in the area of the environment where the combined influence of all environmental phenomena allow for a sustainable metabolism.</p>", "links"=>[], "tags"=>["metabolism-based"], "article_id"=>485563, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g014", "stats"=>{"downloads"=>1, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_6_Metabolism_based_behavior_does_not_respond_directly_to_environmental_features_but_rather_to_their_combined_effect_upon_metabolism_/485563", "title"=>"Experiment 6: Metabolism-based behavior does not respond directly to environmental features, but rather to their combined effect upon metabolism.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:04:26"}
  • {"files"=>["https://ndownloader.figshare.com/files/814741"], "description"=>"<p>Placed in an environment with two sets of metabolizable resources ( located in the lower-right and located in the upper-right), the simulated bacteria move from an initial, even-distribution to the areas higher in concentration of either resource-pair. The insertion of (a chemical that inhibits the metabolism of ) causes the simulated bacteria to cease chemotaxis towards the no-longer metabolizable resources without influencing chemotaxis to the other attractants. Histogram A illustrates average distance to the non-inhibited () source. Histogram B shows the distance from the inhibited source (). As before, data is taken from 10 trials of 100 bacteria each. Plot C illustrates the spatial distribution of simulated bacteria in a typical trial.</p>", "links"=>[], "tags"=>["inhibition", "metabolic", "pathway", "inhibits", "chemotaxis"], "article_id"=>485115, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g009", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Experiment_3_Inhibition_of_a_metabolic_pathway_inhibits_chemotaxis_only_to_the_relevant_resources_/485115", "title"=>"Experiment 3: Inhibition of a metabolic pathway inhibits chemotaxis only to the relevant resources.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:02:04"}
  • {"files"=>["https://ndownloader.figshare.com/files/815243"], "description"=>"<p>Full list of the differential equations that influence the change in chemical concentrations. Includes all chemical pathways used in the experiments. Parameters and represent the reaction rate constants for the th reaction in the forward or backward direction. Those constants with ‘degK’ subscripts represent the rate of degradation of chemical K and constants with ‘’ subscripts represent the concentration of resource diffusing into the system from the environment at position . <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1001004#pcbi-1001004-t001\" target=\"_blank\">Table 1</a> indicates the values of the constants.</p>", "links"=>[], "tags"=>["cell biology/chemical biology of the cell", "computational biology/metabolic networks", "computational biology/systems biology", "microbiology/microbial physiology and metabolism"], "article_id"=>485612, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.t002", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Differential_equations_/485612", "title"=>"Differential equations.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-02-20 22:04:42"}
  • {"files"=>["https://ndownloader.figshare.com/files/814096"], "description"=>"<p>CW rotation results in a random re-orientation for the bacterium, but CCW rotation of flagellar motors produces an approximately straight-line motion.</p>", "links"=>[], "tags"=>["modes"], "article_id"=>484465, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g001", "stats"=>{"downloads"=>2, "page_views"=>149, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Tumbling_and_running_modes_associated_with_Escherichia_coli_and_Salmonella_typhimurium_/484465", "title"=>"Tumbling and running modes associated with <i>Escherichia coli</i> and <i>Salmonella typhimurium</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 21:58:38"}
  • {"files"=>["https://ndownloader.figshare.com/files/814169"], "description"=>"<p>Arrows indicate <i>only</i> short-term dynamical influence between processes.</p>", "links"=>[], "tags"=>["metabolism-independent", "metabolism-based"], "article_id"=>484539, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g002", "stats"=>{"downloads"=>1, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Differences_between_metabolism_dependent_metabolism_independent_and_metabolism_based_chemotaxis_/484539", "title"=>"Differences between metabolism-dependent, metabolism-independent and metabolism-based chemotaxis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 21:59:05"}
  • {"files"=>["https://ndownloader.figshare.com/files/815000"], "description"=>"<p>Diagram of the metabolic reactions for Experiment 4.</p>", "links"=>[], "tags"=>["metabolic", "reactions"], "article_id"=>485372, "categories"=>["Computational Biology", "Cell Biology", "Microbiology", "Medicine"], "users"=>["Matthew D. Egbert", "Xabier E. Barandiaran", "Ezequiel A. Di Paolo"], "doi"=>"https://dx.doi.org/10.1371/journal.pcbi.1001004.g012", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Diagram_of_the_metabolic_reactions_for_Experiment_4_/485372", "title"=>"Diagram of the metabolic reactions for Experiment 4.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-20 22:03:29"}

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  • {"unique-ip"=>"7", "full-text"=>"5", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"4", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"8"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"2", "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"=>"12", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"7", "full-text"=>"7", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"4", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"11"}
  • {"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"=>"12"}
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  • {"unique-ip"=>"8", "full-text"=>"4", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"4", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"2"}
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  • {"unique-ip"=>"5", "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"=>"5"}
  • {"unique-ip"=>"11", "full-text"=>"11", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2017", "month"=>"6"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"7"}
  • {"unique-ip"=>"8", "full-text"=>"7", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"8"}
  • {"unique-ip"=>"3", "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"=>"9"}
  • {"unique-ip"=>"7", "full-text"=>"5", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
  • {"unique-ip"=>"8", "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"=>"11"}
  • {"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"=>"12"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"6", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"1"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"3"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"1"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"6", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"5"}
  • {"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"=>"6"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"8", "full-text"=>"4", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"5", "full-text"=>"7", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"11"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"2"}
  • {"unique-ip"=>"8", "full-text"=>"7", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
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  • {"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"=>"5"}
  • {"unique-ip"=>"11", "full-text"=>"11", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"5", "full-text"=>"6", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
  • {"unique-ip"=>"9", "full-text"=>"9", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"10"}
  • {"unique-ip"=>"15", "full-text"=>"9", "pdf"=>"9", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"12"}
  • {"unique-ip"=>"14", "full-text"=>"12", "pdf"=>"6", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"2", "year"=>"2020", "month"=>"2"}
  • {"unique-ip"=>"10", "full-text"=>"10", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"3"}
  • {"unique-ip"=>"5", "full-text"=>"8", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"4"}
  • {"unique-ip"=>"15", "full-text"=>"17", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"5"}
  • {"unique-ip"=>"10", "full-text"=>"11", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"6"}
  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"7"}
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Relative Metric

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