Helper Response to Experimentally Manipulated Predation Risk in the Cooperatively Breeding Cichlid Neolamprologus pulcher
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{"title"=>"Helper response to experimentally manipulated predation risk in the cooperatively breeding cichlid Neolamprologus pulcher", "type"=>"journal", "authors"=>[{"first_name"=>"Dik", "last_name"=>"Heg", "scopus_author_id"=>"6701630557"}, {"first_name"=>"Michael", "last_name"=>"Taborsky", "scopus_author_id"=>"7004445566"}], "year"=>2010, "source"=>"PLoS ONE", "identifiers"=>{"scopus"=>"2-s2.0-77956276372", "sgr"=>"77956276372", "issn"=>"19326203", "doi"=>"10.1371/journal.pone.0010784", "pmid"=>"20520816", "isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)", "pui"=>"359515753"}, "id"=>"4a7067b6-b12c-3d80-9799-5fd65d4af289", "abstract"=>"Background\\nWe manipulated predation risk in a field experiment with the cooperatively breeding cichlid Neolamprologus pulcher by releasing no predator, a medium- or a large-sized fish predator inside underwater cages enclosing two to three natural groups. We assessed whether helpers changed their helping behaviour, and whether within-group conflict changed, depending on these treatments, testing three hypotheses: ‘pay-to-stay’ PS, ‘risk avoidance’ RA, or (future) reproductive benefits RB. We also assessed whether helper food intake was reduced under risk, because this might reduce investments in other behaviours to save energy.\\n\\nMethodology/Principal Findings\\nMedium and large helpers fed less under predation risk. Despite this effect helpers invested more in territory defence, but not territory maintenance, under the risk of predation (supporting PS). Experimentally covering only the breeding shelter with sand induced more helper digging under predation risk compared to the control treatment (supporting PS). Aggression towards the introduced predator did not differ between the two predator treatments and increased with group member size and group size (supporting PS and RA). Large helpers increased their help ratio (helping effort/breeder aggression received, ‘punishment’ by the dominant pair in the group) in the predation treatments compared to the control treatment, suggesting they were more willing to PS. Medium helpers did not show such effects. Large helpers also showed a higher submission ratio (submission/ breeder aggression received) in all treatments, compared to the medium helpers (supporting PS).\\n\\nConclusions/Significance\\nWe conclude that predation risk reduces helper food intake, but despite this effect, helpers were more willing to support the breeders, supporting PS. Effects of breeder punishment suggests that PS might be more important for large compared to the medium helpers. Evidence for RA was also detected. Finally, the results were inconsistent with RB.", "link"=>"http://www.mendeley.com/research/helper-response-experimentally-manipulated-predation-risk-cooperatively-breeding-cichlid-neolamprolo", "reader_count"=>56, "reader_count_by_academic_status"=>{"Student > Doctoral Student"=>3, "Researcher"=>12, "Student > Ph. D. Student"=>23, "Student > Postgraduate"=>1, "Student > Master"=>7, "Student > Bachelor"=>6, "Lecturer"=>1, "Professor"=>3}, "reader_count_by_user_role"=>{"Student > Doctoral Student"=>3, "Researcher"=>12, "Student > Ph. D. Student"=>23, "Student > Postgraduate"=>1, "Student > Master"=>7, "Student > Bachelor"=>6, "Lecturer"=>1, "Professor"=>3}, "reader_count_by_subject_area"=>{"Unspecified"=>1, "Environmental Science"=>2, "Agricultural and Biological Sciences"=>52, "Psychology"=>1}, "reader_count_by_subdiscipline"=>{"Psychology"=>{"Psychology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>52}, "Unspecified"=>{"Unspecified"=>1}, "Environmental Science"=>{"Environmental Science"=>2}}, "reader_count_by_country"=>{"Netherlands"=>1, "Romania"=>1, "United States"=>1, "Japan"=>1, "United Kingdom"=>1, "Switzerland"=>2, "Portugal"=>1, "Germany"=>2, "Spain"=>1}, "group_count"=>1}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/847875"], "description"=>"a<p>If helpers showed any predator defence in the predator treatments.</p>b<p>Help or submission as ratio of breeder punishment.</p>c<p>No effect in the focal observations, but effect in digging experiment.</p>d<p>Helpers joined defence against the introduced predator, but also clear evidence for risk avoidance due to aggression declining with <i>N. pulcher</i> body size.</p>e<p>No clear predictions here, but there was a correlation between helper submissiveness and breeder punishment (supporting risk avoidance), which was not matched by an increase in the submission ratio in the predator treatments compared to the control treatment (contra risk avoidance).</p>", "links"=>[], "tags"=>["treatments", "hypotheses", "mentioned", "observed", "differences"], "article_id"=>518334, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.t003", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Expected_effects_of_the_predator_treatments_under_the_three_hypotheses_mentioned_in_the_introduction_and_the_observed_differences_predator_treatments_lt_control_treatment_0_predator_treatments_8202_8202_control_treatment_predator_treatments_gt_control_tr/518334", "title"=>"Expected effects of the predator treatments under the three hypotheses mentioned in the introduction and the observed differences (-: predator treatments < control treatment, 0: predator treatments  =  control treatment, +: predator treatments > control treatment).", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-05-26 02:18:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/847370"], "description"=>"<p>Depicted are means ± s.e.m. per 15 minutes observation time, except (A) per minute not hiding, of behaviour depending on the treatments (white circles: control; black triangles: medium predator; black squares: large predator) and helper size (small symbols and thin lines: medium helpers; large symbols and bold lines: large helpers). For statistics see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0010784#pone.0010784.s001\" target=\"_blank\">Table S1</a>. (A) Feeding rate per minute not hiding, (B) total food intake, (C) digging frequency, (D) territory defence frequency (excluding against introduced predator), (E) within-group conflicts, (F) within-group social contacts. Sample sizes are <i>n</i> = 42 for each symbol, except for medium helpers large predator treatment (<i>n</i> = 41 due to one sample lost).</p>", "links"=>[], "tags"=>["subordinate", "behaviour"], "article_id"=>517832, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.g001", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Focal_subordinate_behaviour_depending_on_the_predator_treatments_/517832", "title"=>"Focal subordinate behaviour depending on the predator treatments.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-26 02:10:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/421794"], "description"=>"<div><h3>Background</h3><p>We manipulated predation risk in a field experiment with the cooperatively breeding cichlid <em>Neolamprologus pulcher</em> by releasing no predator, a medium- or a large-sized fish predator inside underwater cages enclosing two to three natural groups. We assessed whether helpers changed their helping behaviour, and whether within-group conflict changed, depending on these treatments, testing three hypotheses: ‘pay-to-stay’ PS, ‘risk avoidance’ RA, or (future) reproductive benefits RB. We also assessed whether helper food intake was reduced under risk, because this might reduce investments in other behaviours to save energy.</p><h3>Methodology/Principal Findings</h3><p>Medium and large helpers fed less under predation risk. Despite this effect helpers invested more in territory defence, but not territory maintenance, under the risk of predation (supporting PS). Experimentally covering only the breeding shelter with sand induced more helper digging under predation risk compared to the control treatment (supporting PS). Aggression towards the introduced predator did not differ between the two predator treatments and increased with group member size and group size (supporting PS and RA). Large helpers increased their help ratio (helping effort/breeder aggression received, ‘punishment’ by the dominant pair in the group) in the predation treatments compared to the control treatment, suggesting they were more willing to PS. Medium helpers did not show such effects. Large helpers also showed a higher submission ratio (submission/ breeder aggression received) in all treatments, compared to the medium helpers (supporting PS).</p><h3>Conclusions/Significance</h3><p>We conclude that predation risk reduces helper food intake, but despite this effect, helpers were more willing to support the breeders, supporting PS. Effects of breeder punishment suggests that PS might be more important for large compared to the medium helpers. Evidence for RA was also detected. Finally, the results were inconsistent with RB.</p></div>", "links"=>[], "tags"=>["helper", "experimentally", "manipulated", "predation", "cooperatively", "breeding", "cichlid"], "article_id"=>143299, "categories"=>["Evolutionary Biology", "Marine Biology", "Cancer"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784", "stats"=>{"downloads"=>4, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Helper_Response_to_Experimentally_Manipulated_Predation_Risk_in_the_Cooperatively_Breeding_Cichlid_Neolamprologus_pulcher_/143299", "title"=>"Helper Response to Experimentally Manipulated Predation Risk in the Cooperatively Breeding Cichlid <em>Neolamprologus pulcher</em>", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-05-26 00:54:59"}
  • {"files"=>["https://ndownloader.figshare.com/files/847917"], "description"=>"<p>Note that for practical reasons effort was measured for all individuals per status lumped, so the number of individuals observed per status was entered as a covariate to arrive at per capita estimates.</p><p>Results were corrected for random group effects (repeated measures of 24 groups for digging and 33 groups for attacks, entered as subjects in the GEEs), the scaling parameter was adjusted using the deviance method in each model.</p>a<p>Group member status was divided in five classes (<i>n</i> = 24 in each case): small helper (15.5–25.0 mm SL, mean individuals per group ± s.d., range: 2.58±1.35, 0–6), medium helper (25.5–35 mm SL, 2.33±1.58, 1–8), large helper (>35 mm SL, 2.33±1.17, 1–6), breeder female (1.04±0.36, 0–2) or breeder male (0.96±0.20, 0–1), so in total 222 individuals were observed. Small helpers were missing for one group with a medium predator, and both the breeder female and breeder male were missing from another group with a medium predator.</p>b<p>Group member status was divided in five classes (<i>n</i> = 33 in each case): small helper (15.5–25.0 mm SL, mean individuals per group ± s.d., range: 2.74±2.03, 1–10), medium helper (25.5–35 mm SL, 2.45±1.58, 1–8), large helper (>35 mm SL, 4.88±2.83, 2–17), breeder female (1.09±0.29, 1–2) or breeder male (1.00±0.00, 1–1), and since these groups were observed multiply we had 803 cases in total. However, since small helpers were never seen to attack the introduced predators, they were omitted from the GEE and this reduced the sample size to 664 (<i>n</i> = 33 groups with 8 to 36 measurements per group).</p>", "links"=>[], "tags"=>["attacks", "introduced", "poisson", "gees", "digging", "carrying", "breeding", "was", "covered"], "article_id"=>518366, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.t001", "stats"=>{"downloads"=>6, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Digging_experiment_and_attacks_on_the_introduced_predator_results_of_two_poisson_GEEs_with_log_link_testing_for_fixed_effects_of_the_treatment_status_and_their_interaction_on_the_digging_plus_sand_carrying_effort_after_the_breeding_shelter_was_covered_wi/518366", "title"=>"Digging experiment and attacks on the introduced predator: results of two poisson GEEs with log-link, testing for fixed effects of the treatment, status and their interaction on the digging plus sand carrying effort after the breeding shelter was covered with sand (<i>n</i> = 117) and attacks on the introduced predator (<i>n</i> = 664) separately.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-05-26 02:19:26"}
  • {"files"=>["https://ndownloader.figshare.com/files/847631"], "description"=>"<p>(A) Shows the introduced predator on the left (<i>L. elongatus</i>) and defending group members of <i>N. pulcher</i> on the right. Depicted are means ± s.e.m. of aggression and sample sizes (number of groups) for the five different types of groups members in the (B) medium and (C) large predator treatment (Sh: Small helpers, Mh: Medium helpers, Lh: Large helpers, Bf: breeding females, Bm: breeding males). Note that small helpers were never seen to attack the medium and large predators (see text). For statistics see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0010784#pone-0010784-t001\" target=\"_blank\">Table 1</a>.</p>", "links"=>[], "tags"=>["capita", "introduced"], "article_id"=>518095, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.g004", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_per_capita_frequency_of_aggression_against_the_introduced_predator_/518095", "title"=>"The per capita frequency of aggression against the introduced predator.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-26 02:14:55"}
  • {"files"=>["https://ndownloader.figshare.com/files/847548"], "description"=>"<p>Effect of covering the breeding shelter with sand on subsequent digging and carrying sand behaviour from this shelter by the different group members in the three predator treatments. Depicted are means ± s.e.m. and sample sizes (number of groups). For statistics see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0010784#pone-0010784-t001\" target=\"_blank\">Table 1</a>.</p>", "links"=>[], "tags"=>["evolutionary biology/animal behavior", "evolutionary biology/evolutionary ecology", "marine and aquatic sciences/ecology", "marine and aquatic sciences/evolutionary biology"], "article_id"=>518013, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.g003", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Digging_experiment_/518013", "title"=>"Digging experiment.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-26 02:13:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/847754"], "description"=>"<p>The ratio of (A) helping effort and (B) submission shown to the breeders, per received aggression from the breeders depending on the predator treatments (white circles: control; black triangles: medium predator; black squares: large predator) and helper size (small symbols and thin lines: medium helpers; large symbols and bold lines: large helpers). Depicted are means ± s.e.m (ratios =  [frequency helping or submission square-root +3/8 transformed]/[frequency breeder aggression received square-root +3/8 transformed]) with sample sizes (number of observations). For statistics see text. Note that the sample size for medium helpers in the large predator treatment was <i>n</i> = 40 due to one sample lost and one sample the helper was hiding 100% of the time (gives 42–2 = 40).</p>", "links"=>[], "tags"=>["evolutionary biology/animal behavior", "evolutionary biology/evolutionary ecology", "marine and aquatic sciences/ecology", "marine and aquatic sciences/evolutionary biology"], "article_id"=>518213, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.g005", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Helper_breeder_conflict_/518213", "title"=>"Helper-breeder conflict.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-26 02:16:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/847835"], "description"=>"<p>In brackets sample sizes.</p><p>*<i>p</i><0.05,</p><p>**<i>p</i><0.001.</p>", "links"=>[], "tags"=>["spearman", "correlations", "capita", "predators", "members", "adults", "breeders", "treatments"], "article_id"=>518287, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.t002", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Results_of_Spearman_Rank_Correlations_between_the_per_capita_frequency_of_aggression_against_the_predators_by_the_different_group_members_and_the_number_of_adults_living_in_the_group_number_of_breeders_and_large_helpers_for_the_medium_above_diagonal_and_/518287", "title"=>"Results of Spearman Rank Correlations between the per capita frequency of aggression against the predators by the different group members and the number of adults living in the group (number of breeders and large helpers), for the medium (above diagonal) and large predator treatments (below diagonal), separately.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-05-26 02:18:07"}
  • {"files"=>["https://ndownloader.figshare.com/files/847431"], "description"=>"<p>Intake rate increased with average distance moved from protective shelter in both (A) large helpers and (B) medium helpers. Depicted are hyperbolic curve fits (distance/[a+b x distance]) from non-linear regressions for large helpers (<i>n</i> = 126, coefficients ± s.e.): a = 1.1572±0.2642, b = 0.0257±0.0064 (<i>F</i><sub>2,124</sub> = 229.7, <i>p</i><0.001, <i>R</i><sup>2</sup> = 0.23); and for medium helpers (<i>n</i> = 125): a = 0.4575±0.0910, b = 0.0342±0.0042 (<i>F</i><sub>2,123</sub> = 226.6, <i>p</i><0.001, <i>R</i><sup>2</sup> = 0.30).</p>", "links"=>[], "tags"=>["subordinate", "feeding", "was", "spacing"], "article_id"=>517887, "categories"=>["Evolutionary Biology", "Marine Biology", "Infectious Diseases"], "users"=>["Dik Heg", "Michael Taborsky"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0010784.g002", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Focal_subordinate_feeding_rate_was_closely_related_to_spacing_behaviour_/517887", "title"=>"Focal subordinate feeding rate was closely related to spacing behaviour.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-26 02:11:27"}

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  • {"unique-ip"=>"4", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"8"}
  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"9"}
  • {"unique-ip"=>"10", "full-text"=>"9", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2015", "month"=>"10"}
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  • {"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"=>"2016", "month"=>"2"}
  • {"unique-ip"=>"12", "full-text"=>"5", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2014", "month"=>"11"}
  • {"unique-ip"=>"7", "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"=>"12"}
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  • {"unique-ip"=>"8", "full-text"=>"7", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2015", "month"=>"11"}
  • {"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"=>"2015", "month"=>"12"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"1"}
  • {"unique-ip"=>"2", "full-text"=>"0", "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"=>"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"=>"5"}
  • {"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"=>"6"}
  • {"unique-ip"=>"1", "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"=>"7"}
  • {"unique-ip"=>"3", "full-text"=>"7", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"8"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"4", "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"=>"11"}
  • {"unique-ip"=>"9", "full-text"=>"8", "pdf"=>"2", "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"=>"2", "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"=>"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"=>"3"}
  • {"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"=>"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"=>"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"=>"8"}
  • {"unique-ip"=>"3", "full-text"=>"5", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"9"}
  • {"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"=>"10"}
  • {"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"=>"2017", "month"=>"11"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"12"}
  • {"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"=>"2018", "month"=>"1"}
  • {"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"=>"3"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"1"}
  • {"unique-ip"=>"9", "full-text"=>"9", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"5"}
  • {"unique-ip"=>"4", "full-text"=>"6", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"6"}
  • {"unique-ip"=>"3", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"5", "full-text"=>"2", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"3", "full-text"=>"4", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"9", "full-text"=>"9", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"11"}
  • {"unique-ip"=>"10", "full-text"=>"13", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
  • {"unique-ip"=>"10", "full-text"=>"6", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"4", "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"=>"3", "full-text"=>"5", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"3"}
  • {"unique-ip"=>"7", "full-text"=>"10", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"4"}
  • {"unique-ip"=>"4", "full-text"=>"4", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"5"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"7", "full-text"=>"11", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
  • {"unique-ip"=>"6", "full-text"=>"5", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2019", "month"=>"10"}
  • {"unique-ip"=>"5", "full-text"=>"3", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"12"}
  • {"unique-ip"=>"8", "full-text"=>"10", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"2"}
  • {"unique-ip"=>"10", "full-text"=>"11", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"3"}
  • {"unique-ip"=>"13", "full-text"=>"16", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"4"}
  • {"unique-ip"=>"6", "full-text"=>"7", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"5"}
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Relative Metric

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