Collective Defense of Aphis nerii and Uroleucon hypochoeridis (Homoptera, Aphididae) against Natural Enemies
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{"title"=>"Collective defense of aphis nerii and uroleucon hypochoeridis (homoptera, aphididae) against natural enemies", "type"=>"journal", "authors"=>[{"first_name"=>"Manfred", "last_name"=>"Hartbauer", "scopus_author_id"=>"8635880600"}], "year"=>2010, "source"=>"PLoS ONE", "identifiers"=>{"issn"=>"19326203", "pui"=>"359495613", "pmid"=>"20454683", "isbn"=>"1932-6203", "doi"=>"10.1371/journal.pone.0010417", "sgr"=>"77956446094", "scopus"=>"2-s2.0-77956446094"}, "id"=>"2dc31235-3c9f-3b15-a121-bebe8320112c", "abstract"=>"The prevalent way aphids accomplish colony defense against natural enemies is a mutualistic relationship with ants or the occurrence of a specialised soldier caste typical for eusocial aphids, or even both. Despite a group-living life style of those aphid species lacking these defense lines, communal defense against natural predators has not yet been observed there. Individuals of Aphis nerii (Oleander aphid) and Uroleucon hypochoeridis, an aphid species feeding on Hypochoeris radicata (hairy cat's ear), show a behavioral response to visual stimulation in the form of spinning or twitching, which is often accompanied by coordinated kicks executed with hind legs. Interestingly, this behaviour is highly synchronized among members of a colony and repetitive visual stimulation caused strong habituation. Observations of natural aphid colonies revealed that a collective twitching and kicking response (CTKR) was frequently evoked during oviposition attempts of the parasitoid wasp Aphidius colemani and during attacks of aphidophagous larvae. CTKR effectively interrupted oviposition attempts of this parasitoid wasp and even repelled this parasitoid from colonies after evoking consecutive CTKRs. In contrast, solitary feeding A. nerii individuals were not able to successfully repel this parasitoid wasp. In addition, CTKR was also evoked through gentle substrate vibrations. Laser vibrometry of the substrate revealed twitching-associated vibrations that form a train of sharp acceleration peaks in the course of a CTKR. This suggests that visual signals in combination with twitching-related substrate vibrations may play an important role in synchronising defense among members of a colony. In both aphid species collective defense in encounters with different natural enemies was executed in a stereotypical way and was similar to CTKR evoked through visual stimulation. This cooperative defense behavior provides an example of a surprising sociality that can be found in some aphid species that are not expected to be social at all.", "link"=>"http://www.mendeley.com/research/collective-defense-aphis-nerii-uroleucon-hypochoeridis-homoptera-aphididae-against-natural-enemies", "reader_count"=>59, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>3, "Librarian"=>1, "Researcher"=>15, "Student > Doctoral Student"=>6, "Student > Ph. D. Student"=>14, "Student > Postgraduate"=>1, "Student > Master"=>7, "Other"=>3, "Student > Bachelor"=>3, "Lecturer > Senior Lecturer"=>2, "Professor"=>2}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>3, "Librarian"=>1, "Researcher"=>15, "Student > Doctoral Student"=>6, "Student > Ph. D. Student"=>14, "Student > Postgraduate"=>1, "Student > Master"=>7, "Other"=>3, "Student > Bachelor"=>3, "Lecturer > Senior Lecturer"=>2, "Professor"=>2}, "reader_count_by_subject_area"=>{"Engineering"=>2, "Unspecified"=>5, "Environmental Science"=>4, "Agricultural and Biological Sciences"=>38, "Neuroscience"=>1, "Psychology"=>2, "Social Sciences"=>1, "Computer Science"=>6}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>2}, "Neuroscience"=>{"Neuroscience"=>1}, "Social Sciences"=>{"Social Sciences"=>1}, "Psychology"=>{"Psychology"=>2}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>38}, "Computer Science"=>{"Computer Science"=>6}, "Unspecified"=>{"Unspecified"=>5}, "Environmental Science"=>{"Environmental Science"=>4}}, "reader_count_by_country"=>{"Colombia"=>1, "Canada"=>1, "United States"=>3, "Mexico"=>1, "France"=>2, "Switzerland"=>1, "Portugal"=>1, "Germany"=>1, "India"=>1}, "group_count"=>4}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/851878"], "description"=>"<p><i>A. nerii</i> colonies feeding on Oleander (<b>A</b>–<b>C</b>) and <i>U. hypochoeridis</i> colonies feeding on <i>Hypochoeris radicata</i> (<b>D–F</b>). CTKR of <i>A. nerii</i> colonies is restricted to those colonies consisting of bucking individuals (a lift abdomen) and some space left for twitching (B). The colony in A is not responding to a rapidly approaching object like a ball pen (diameter  = 12 mm, tip size  = 1 mm). <b>C </b><i>Aphidius colemani</i> after less successful oviposition attempts. <b>E</b> Members of an <i>U. hypochoeridis</i> colony in defense posture. <b>F</b> An oviposition attempt of <i>Aphidius colemani</i> before this parasitoid was repelled from an <i>U. hypochoeridis</i> colony through consecutive CTKRs.</p>", "links"=>[], "tags"=>["aphid", "colonies", "parasitoid", "wasp"], "article_id"=>522340, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g002", "stats"=>{"downloads"=>3, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Pictures_of_aphid_colonies_and_the_parasitoid_wasp_Aphidius_colemani_/522340", "title"=>"Pictures of aphid colonies and the parasitoid wasp <i>Aphidius colemani</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:39:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/852338"], "description"=>"<p>Quickly moving back and forth the tip of a blue ball pen (see S1A) resulted in a habituation of CTKR of <i>A. nerii</i> (A) and <i>U. hypochoeridis</i> (B) colonies. Stimulus directions are indicated by double-arrows in the pictures next to A and B. The stimulus in A was either presented from the right side (blue curve) or from top (red curve). Introducing a pause of 15 s in <i>A. nerii</i> (last dot in the red curve in A) and a pause of 60 s in <i>U. hypochoeridis</i> restored CTKR (red curve in B). <b>C</b> Time course of CTKR evoked by substrate vibrations that were induced by gently tipping on the stalk 10 cm below the colonized leaf (ellipse in the picture next to C). Vibratory stimulations were separated by pauses of 30 s. Curves in C are aligned to the onset of the stimulus.</p>", "links"=>[], "tags"=>["ctkr", "ctkrs", "evoked", "substrate"], "article_id"=>522793, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g005", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Habituation_of_CTKR_and_CTKRs_evoked_through_substrate_vibrations_/522793", "title"=>"Habituation of CTKR and CTKRs evoked through substrate vibrations.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:46:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/423940", "https://ndownloader.figshare.com/files/423963", "https://ndownloader.figshare.com/files/423980", "https://ndownloader.figshare.com/files/424007", "https://ndownloader.figshare.com/files/424041", "https://ndownloader.figshare.com/files/424078", "https://ndownloader.figshare.com/files/424122", "https://ndownloader.figshare.com/files/424159", "https://ndownloader.figshare.com/files/424214"], "description"=>"<div><p>The prevalent way aphids accomplish colony defense against natural enemies is a mutualistic relationship with ants or the occurrence of a specialised soldier caste typcial for eusocial aphids, or even both. Despite a group-living life style of those aphid species lacking these defense lines, communal defense against natural predators has not yet been observed there. Individuals of <em>Aphis nerii</em> (Oleander aphid) and <em>Uroleucon hypochoeridis</em>, an aphid species feeding on <em>Hypochoeris radicata</em> (hairy cat's ear), show a behavioral response to visual stimulation in the form of spinning or twitching, which is often accompanied by coordinated kicks executed with hind legs. Interestingly, this behaviour is highly synchronized among members of a colony and repetitive visual stimulation caused strong habituation. Observations of natural aphid colonies revealed that a collective twitching and kicking response (CTKR) was frequently evoked during oviposition attempts of the parasitoid wasp <em>Aphidius colemani</em> and during attacks of aphidophagous larvae. CTKR effectively interrupted oviposition attempts of this parasitoid wasp and even repelled this parasitoid from colonies after evoking consecutive CTKRs. In contrast, solitary feeding <em>A. nerii</em> individuals were not able to successfully repel this parasitoid wasp. In addition, CTKR was also evoked through gentle substrate vibrations. Laser vibrometry of the substrate revealed twitching-associated vibrations that form a train of sharp acceleration peaks in the course of a CTKR. This suggests that visual signals in combination with twitching-related substrate vibrations may play an important role in synchronising defense among members of a colony. In both aphid species collective defense in encounters with different natural enemies was executed in a stereotypical way and was similar to CTKR evoked through visual stimulation. This cooperative defense behavior provides an example of a surprising sociality that can be found in some aphid species that are not expected to be social at all.</p></div>", "links"=>[], "tags"=>["enemies"], "article_id"=>143692, "categories"=>["Cancer", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0010417.s001", "https://dx.doi.org/10.1371/journal.pone.0010417.s002", "https://dx.doi.org/10.1371/journal.pone.0010417.s003", "https://dx.doi.org/10.1371/journal.pone.0010417.s004", "https://dx.doi.org/10.1371/journal.pone.0010417.s005", "https://dx.doi.org/10.1371/journal.pone.0010417.s006", "https://dx.doi.org/10.1371/journal.pone.0010417.s007", "https://dx.doi.org/10.1371/journal.pone.0010417.s008", "https://dx.doi.org/10.1371/journal.pone.0010417.s009"], "stats"=>{"downloads"=>13, "page_views"=>20, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Collective_Defense_of_Aphis_nerii_and_Uroleucon_hypochoeridis_Homoptera_Aphididae_against_Natural_Enemies/143692", "title"=>"Collective Defense of <em>Aphis nerii</em> and <em>Uroleucon hypochoeridis</em> (Homoptera, Aphididae) against Natural Enemies", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2010-04-29 01:01:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/852079"], "description"=>"<p>Time course of simultaneous twitching observed in <i>A. nerii</i> colonies (A and B) and <i>U. hypochoeridis</i> colonies (C and D). Repeated stabbing attacks of <i>A. colemani</i> evoked CTKR of the colony shown in the circle in the picture next to A. The time course of simultaneous twitching obtained from 6 stabbing attacks are shown in A. <b>B</b> Time course of CTKR of an <i>A. nerii</i> colony evoked by an approaching ball pen (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0010417#pone-0010417-g002\" target=\"_blank\">Fig. 2A</a>). This visual stimulus was always withdrawn after the first aphids started to twitch (∼5 cm in front of the colony). A pause of about 30 sec separates successive stimulations. <b>C </b><i>A. colemani</i> evoked CTKR of an <i>U. hypochoeridis</i> colony shown in the picture next to C. After the third CTKR this parasitoid was repelled (arrow in C). <b>D</b> CTKR of an <i>U. hypochoeridis</i> colony was evoked by a chop stick with a tip diameter of 3 mm. This stick approached this colony either from the right or from left side. CTKR of this colony was also evoked by an approaching insect (size  = 7 mm). The black dot in the flight track (dotted line) indicates the position when the first aphid started to twitch. Arrows in D indicate time points when the stick was withdrawn and the insect steered away from this colony. Data in all graphs are aligned to the first twitching individual.</p>", "links"=>[], "tags"=>["evoked", "parasitoids"], "article_id"=>522541, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g003", "stats"=>{"downloads"=>1, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_CTKR_evoked_by_parasitoids_and_by_visual_stimulation_/522541", "title"=>"CTKR evoked by parasitoids and by visual stimulation.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:42:21"}
  • {"files"=>["https://ndownloader.figshare.com/files/852558"], "description"=>"<p>Summary statistics of CTKR.</p>", "links"=>[], "tags"=>["ecology", "ecology/behavioral ecology", "ecology/evolutionary ecology"], "article_id"=>523023, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.t001", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Summary_statistics_of_CTKR_/523023", "title"=>"Summary statistics of CTKR.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-04-29 00:50:23"}
  • {"files"=>["https://ndownloader.figshare.com/files/852213"], "description"=>"<p>In <i>U. hypochoeridis</i> (A and B) colonies and in <i>A. nerii</i> (C and D) colonies CTKRs were evoked in the course of attacks of ladybird larvae (A), hemerobiid larvae (B and C) and syrphid larvae (D). Curves represent the time course of simultaneously twitching individuals belonging to those aphid colonies that are shown in the pictures next to graphs (ellipse in C and D). Arrows point to aphidophagous larvae. Curves in graphs are aligned to the first twitching individual.</p>", "links"=>[], "tags"=>["evoked", "aphidophagous"], "article_id"=>522677, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g004", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_CTKR_evoked_by_aphidophagous_larvae_/522677", "title"=>"CTKR evoked by aphidophagous larvae.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:44:37"}
  • {"files"=>["https://ndownloader.figshare.com/files/852460"], "description"=>"<p>Lower traces in A and B represent the velocity of substrate vibrations associated with either solitary or collective twitching. Laser vibrometer focus points are labeled as L. <b>A</b> Laser vibrometry of twitching <i>A. nerii</i> individuals (arrows) and small <i>A. nerii</i> colonies (circles) feeding on an Oleander twig. Numbers below the velocity signal in A relate to labels shown in the picture above. <b>B</b> Substrate velocity generated by twitching individuals belonging to a colony of <i>U. hypochoeridis</i> dispersed on three terminal branches of a host plant (<i>Hypochoeris radicata</i>). The acceleration of the velocity signal is shown below sonograms, which were generated by use of a FFT size of 1024 points (sampling rate: 48 kHz). Twitching was evoked by visual stimulation. <b>C</b> The velocity signal marked with an asterisk in A. <b>D</b> The velocity signal marked with an asterisk in B.</p>", "links"=>[], "tags"=>["vibrometry"], "article_id"=>522921, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g006", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_vibrometry_of_CTKR_/522921", "title"=>"Laser vibrometry of CTKR.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:48:41"}
  • {"files"=>["https://ndownloader.figshare.com/files/851801"], "description"=>"<p>Spinning of <i>A. nerii</i> individuals (A, B) is executed in a way so that the tip of the abdomen follows an almost circular path (dotted lines in A). <b>B</b> A visual stimulus presented from the right side (arrow in B) results in a circular movement of the tip of the abdomen and a similar circular movement of the right hind leg. <b>C</b> Resting position of <i>U. hypochoeridis</i>. <b>D</b> and <b>E</b> Successive steps sketching twitching <i>U. hypochoeridis</i> aphids when the stimulus (arrow) is presented on the right side. <b>F</b> Kicks executed by <i>U. hypochoeridis</i> when an object approaches from behind.</p>", "links"=>[], "tags"=>["solitary", "reactions"], "article_id"=>522265, "categories"=>["Infectious Diseases", "Ecology"], "users"=>["Manfred Hartbauer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010417.g001", "stats"=>{"downloads"=>0, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Sketch_of_solitary_defense_reactions_in_A_nerii_and_U_hypochoeridis_aphids_/522265", "title"=>"Sketch of solitary defense reactions in <i>A. nerii</i> and <i>U. hypochoeridis</i> aphids.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-04-29 00:37:45"}

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  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2015", "month"=>"9"}
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  • {"unique-ip"=>"8", "full-text"=>"9", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"6", "supp-data"=>"10", "cited-by"=>"1", "year"=>"2014", "month"=>"10"}
  • {"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"=>"2016", "month"=>"2"}
  • {"unique-ip"=>"6", "full-text"=>"4", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2014", "month"=>"11"}
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  • {"unique-ip"=>"10", "full-text"=>"9", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"12"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"4", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"1"}
  • {"unique-ip"=>"5", "full-text"=>"3", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"3"}
  • {"unique-ip"=>"4", "full-text"=>"4", "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"=>"7", "full-text"=>"8", "pdf"=>"4", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"5"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"6"}
  • {"unique-ip"=>"3", "full-text"=>"1", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"7"}
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  • {"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"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"6", "full-text"=>"3", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"5", "supp-data"=>"18", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"1", "full-text"=>"0", "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"=>"4", "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"=>"12"}
  • {"unique-ip"=>"5", "full-text"=>"6", "pdf"=>"0", "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"=>"0", "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"=>"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"=>"9"}
  • {"unique-ip"=>"8", "full-text"=>"11", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
  • {"unique-ip"=>"9", "full-text"=>"8", "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"=>"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"=>"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"=>"2018", "month"=>"1"}
  • {"unique-ip"=>"14", "full-text"=>"14", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"15", "cited-by"=>"0", "year"=>"2018", "month"=>"3"}
  • {"unique-ip"=>"20", "full-text"=>"19", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"1"}
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  • {"unique-ip"=>"11", "full-text"=>"12", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"15", "full-text"=>"9", "pdf"=>"11", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"6"}
  • {"unique-ip"=>"22", "full-text"=>"16", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"4", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"17", "full-text"=>"18", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"22", "full-text"=>"29", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"11"}
  • {"unique-ip"=>"18", "full-text"=>"28", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"13", "full-text"=>"12", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"27", "full-text"=>"32", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"3", "cited-by"=>"0", "year"=>"2019", "month"=>"2"}
  • {"unique-ip"=>"19", "full-text"=>"22", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
  • {"unique-ip"=>"16", "full-text"=>"20", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"4"}
  • {"unique-ip"=>"20", "full-text"=>"23", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}
  • {"unique-ip"=>"12", "full-text"=>"11", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"6", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"15", "full-text"=>"16", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
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Relative Metric

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