Stoichiometry of HLA Class II-Invariant Chain Oligomers
Publication Date
February 22, 2011
Journal
PLOS ONE
Authors
Norbert Koch, Martin Zacharias, Angelika König, Sebastian Temme, et al
Volume
6
Issue
2
Pages
e17257
DOI
https://dx.plos.org/10.1371/journal.pone.0017257
Publisher URL
http://journals.plos.org/plosone/article?id=10.1371%2Fjournal.pone.0017257
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/21364959
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3043101
Europe PMC
http://europepmc.org/abstract/MED/21364959
Web of Science
000287656600037
Scopus
79952052281
Mendeley
http://www.mendeley.com/research/stoichiometry-hla-class-iiinvariant-chain-oligomers
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Mendeley | Further Information

{"title"=>"Stoichiometry of HLA class II-invariant chain oligomers", "type"=>"journal", "authors"=>[{"first_name"=>"Norbert", "last_name"=>"Koch", "scopus_author_id"=>"55352279100"}, {"first_name"=>"Martin", "last_name"=>"Zacharias", "scopus_author_id"=>"7005598310"}, {"first_name"=>"Angelika", "last_name"=>"König", "scopus_author_id"=>"15742438600"}, {"first_name"=>"Sebastian", "last_name"=>"Temme", "scopus_author_id"=>"21935221200"}, {"first_name"=>"Jürgen", "last_name"=>"Neumann", "scopus_author_id"=>"8121968600"}, {"first_name"=>"Sebastian", "last_name"=>"Springer", "scopus_author_id"=>"8773026400"}], "year"=>2011, "source"=>"PLoS ONE", "identifiers"=>{"issn"=>"19326203", "pmid"=>"21364959", "pui"=>"361337207", "scopus"=>"2-s2.0-79952052281", "doi"=>"10.1371/journal.pone.0017257", "isbn"=>"1932-6203 (Electronic) 1932-6203 (Linking)", "sgr"=>"79952052281"}, "id"=>"5320ca16-bdee-3bf1-aaea-fcb02cf79416", "abstract"=>"BACKGROUND: The HLA gene complex encodes three class II isotypes, DR, DQ, and DP. HLA class II molecules are peptide receptors that present antigens for recognition by T lymphocytes. In antigen presenting cells, the assembly of matched α and β subunits to heterodimers is chaperoned by invariant chain (Ii). Ii forms a homotrimer with three binding sites for class II heterodimers. The current model of class II and Ii structure states that three αβ heterodimers bind to an Ii trimer.\\n\\nMETHODOLOGY/PRINCIPAL FINDINGS: [corrected] We have now analyzed the composition and size of the complexes of class II and Ii using epitope tagged class II subunits and density gradient experiments. We show here that class II-Ii oligomers consist of one class II heterodimer associated with one Ii trimer, such that the DR, DQ and DP isotypes are contained within separate complexes with Ii.\\n\\nCONCLUSION/SIGNIFICANCE: We propose a structural model of the class II-Ii oligomer and speculate that the pentameric class II-Ii complex is bent towards the cell membrane, inhibiting the binding of additional class II heterodimers to Ii.", "link"=>"http://www.mendeley.com/research/stoichiometry-hla-class-iiinvariant-chain-oligomers", "reader_count"=>23, "reader_count_by_academic_status"=>{"Researcher"=>8, "Student > Ph. D. Student"=>6, "Student > Postgraduate"=>1, "Student > Master"=>3, "Student > Bachelor"=>3, "Professor"=>2}, "reader_count_by_user_role"=>{"Researcher"=>8, "Student > Ph. D. Student"=>6, "Student > Postgraduate"=>1, "Student > Master"=>3, "Student > Bachelor"=>3, "Professor"=>2}, "reader_count_by_subject_area"=>{"Unspecified"=>1, "Biochemistry, Genetics and Molecular Biology"=>2, "Agricultural and Biological Sciences"=>16, "Medicine and Dentistry"=>2, "Computer Science"=>1, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>2}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>16}, "Computer Science"=>{"Computer Science"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>2}, "Unspecified"=>{"Unspecified"=>1}}, "reader_count_by_country"=>{"Canada"=>1, "Sweden"=>1, "Netherlands"=>1, "Germany"=>1}, "group_count"=>1}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/796739"], "description"=>"<p>A) SDS-PAGE separated Raji cell extracts from fractions 1 to 24 and western blotting for Ii (monoclonal antibody Bu43) (upper panel) results in two peaks (scan lower panel). B) Immunoprecipitation of DR, and SDS-PAGE separation was conducted as in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0017257#pone-0017257-g002\" target=\"_blank\">Fig. 2C</a>, followed by immunoblotting for Ii (monoclonal antibody Bu43, upper panel) and evaluation as a scan (lower panel). The DR-associated Ii appears in lanes 12 to 16. C) DR and Ii oligomers displayed in a sceme. Peptide-loaded DR (pDRαβ) largely co-migrates in the sucrose gradient with the Ii trimer. Complexes containing DR-Ii are separated in fractions 12 to 16.</p>", "links"=>[], "tags"=>["ii", "sucrose", "gradient", "fractions", "delineation", "dr"], "article_id"=>467104, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g003", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Detection_of_Ii_in_sucrose_gradient_fractions_and_delineation_of_DR_and_Ii_complexes_/467104", "title"=>"Detection of Ii in sucrose gradient fractions and delineation of DR and Ii complexes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 01:58:24"}
  • {"files"=>["https://ndownloader.figshare.com/files/797072"], "description"=>"<p>A) MelJuSo cells transfected with DRαV5 cDNA were lysed. Cell lysates and V5-immunoprecipitates from cell lysates were digested with EndoH (E<sub>H</sub>) (lanes 2 and 5), with PNGse F (P<sub>F</sub>) (lanes 3 and 6), or left untreated (Ø) (lanes 1 and 4). SDS-PAGE separated cell lysates were immunoblotted for DRαV5 (monoclonal antibody V5) and V5-immunoprecipitated for DRα (western blotted with DRα monoclonal antibody TAL-1B5). B) Reactivity of monoclonal antibody TAL-1B5 to V5-tagged DRα is abolished. COS-7 cells were transfected with DRαV5 encoding cDNA. Cell lysate was immunoprecipitated with V5 or with TAL-1B5 (DRα) monoclonal antibodies. Immunoprecipitates were separated in lanes 1 to 4, and lane 5 contains lysate of MelJuSo cells. Lanes were western blotted as indicated. C) COS-7 cells were transfected with DRαV5, DRα, DRβ and Ii encoding cDNAs. Cell lysates from transfected (+) and from non-transfected (−) COS cells were immunoprecipitated for V5 (lanes 1 to 6). Immunoprecipitates and cell lysates (lanes 7 to 10) were SDS-PAGE separated and western blotted with mAbs V5 for DRαV5, TAL-1B5 for DRα, LGII-612.14 for DRβ, and Bu43 for Ii.</p>", "links"=>[], "tags"=>["ab-epitope", "tagged", "chains", "transfected", "meljuso"], "article_id"=>467437, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g006", "stats"=>{"downloads"=>1, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Isolation_of_Ab_epitope_tagged_DR_945_chains_from_transfected_MelJuSo_cells_/467437", "title"=>"Isolation of Ab-epitope tagged DRα chains from transfected MelJuSo cells.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:03:57"}
  • {"files"=>["https://ndownloader.figshare.com/files/797150"], "description"=>"<p>Sucrose gradient fractions of Raji cell lysates were subjected to immunoprecipitation with mAb 1a3 against DQ and with DR mAb ISCR3. Fractions 7 and 8 or 14 and 15 were pooled and immunoprecipitated. DQ (lanes 1 and 2) or DR (lanes 3 and 4) were isolated, and western blotted for DRα (upper panel, mAb TAL-1B5) or for DQβ (lower panel, lanes 1 and 2) and for DRβ (lanes 3 and 4) with mAb LGII-612.14. The position of DRα, of DQβ or of DRβ and bands corresponding to immunoglobulin heavy (H) and light (L) chains are labeled by arrows.</p>", "links"=>[], "tags"=>["dq-ii", "complexes", "sucrose", "gradient"], "article_id"=>467515, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g007", "stats"=>{"downloads"=>4, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Isolation_of_DQ_Ii_complexes_from_sucrose_gradient_fractions_/467515", "title"=>"Isolation of DQ-Ii complexes from sucrose gradient fractions.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:05:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/797222"], "description"=>"<p>The composition of DR-Ii complexes in pulse labeled immunoprecipitates from MelJuSo cells, stably transfected with DRβV5 encoding cDNA. COS cells and MelJuSo cells expressing MHCII subunits and Ii were pulse labeled for 10 min with <sup>35</sup>S-methionine. Cells were lysed and immunoprecipitated for Ii (monoclonal antibody Bu45, lane 1), for DRβ (S35, lane 2), for DRα (monoclonal antibody TAL-1B5, lane 3), for DRβV5 (monoclonal antibody V5, lanes 4 and 5), for DRαβ (monoclonal antibody I251SB, lanes 6 and 7) and as a control of reactivity of V5 monoclonal antibody with untransfected MelJuSo cell extract (monoclonal antibody V5, lane 8). Lanes 9 and 10 show molecular weight standards. In lane 11 a DRβV5 immunoprecipitate of MelJuSo cells which were pulse labeled for 10 min with <sup>35</sup>S methionine and subsequently cultured for 40 min with medium containing non-radioactive methionine is shown. Immunoprecipitates from lanes 4 and 5 (left panel) were separated on a 20 cm long SDS PAGE gel to separate DRβV5 and DRα chains (right panel).</p>", "links"=>[], "tags"=>["labeling", "reveals", "ii"], "article_id"=>467591, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g008", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Metabolic_labeling_reveals_that_only_one_DR_946_chain_associated_to_an_Ii_homotrimer_/467591", "title"=>"Metabolic labeling reveals that only one DRβ chain associated to an Ii homotrimer.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:06:31"}
  • {"files"=>["https://ndownloader.figshare.com/files/796872"], "description"=>"<p>DR and Ii oligomers separated by sucrose gradient fractions of Raji cell lysates were crosslinked with DSP. A) Sucrose gradient fractions 7/8 and fractions 14/15 were analysed in the upper or lower panels. Concentration of DSP is shown on top. Untreated or crosslinked samples were separated by SDS-PAGE and western blotted for DRα (monoclonal antibody TAL-1B5, left panels) or for Ii (monoclonal antibody Bu43, right panels). On the right, positions of molecular weight markers are depicted. The dimeric form of Ii (70 kDa), which is seen both in untreated and in crosslinked samples, is produced by oxidation of a free cysteine residue of Ii to an inter-molecular disulfide bond <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0017257#pone.0017257-Koch1\" target=\"_blank\">[24]</a>. B) Fractions 14/15 were crosslinked with 200 µg DSP/ml. Subsequently, crosslinked proteins were subjected to ultra-centrifugation on a sucrose gradient, separated by reducing SDS-PAGE followed by immunoblotting for DRα.</p>", "links"=>[], "tags"=>["crosslinking", "dr", "ii"], "article_id"=>467238, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g004", "stats"=>{"downloads"=>1, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Chemical_crosslinking_of_DR_and_of_Ii_oligomers_/467238", "title"=>"Chemical crosslinking of DR and of Ii oligomers.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:00:38"}
  • {"files"=>["https://ndownloader.figshare.com/files/796452"], "description"=>"<p>Cell lysates were ultra-centrifugated on a 0.4 to 0.8 M sucrose gradient at 100 000 g for 50 h. A) The sucrose concentrations of the fractions were determined after ultracentrifugation by a refractometer (upper panel). In a parallel tube, molecular weight standards were separated (lower panel). The SDS-PAGE separated protein standards were stained with Coomassie blue. The peak positions of the separated molecular weight markers are indicated by arrows. IgG bands (insert) are shown in fractions 14 and 15. B) Twenty four samples, obtained from the fractions of the sucrose gradient of Raji cell lysates were boiled in SDS sample buffer and analysed by SDS-PAGE and subsequent western blotting with monoclonal antibody TAL-1B5 against DRα (upper panel). C) Cell lysates from monocyte-derived human dendritic cells separated by the sucrose gradient were western blotted for DRα. The positions of DRα and of a protein standard (34 kDa) are indicated by arrows. The lower panels of B and C show densitometric scans of the western blots.</p>", "links"=>[], "tags"=>["extracts", "sucrose", "gradient"], "article_id"=>466819, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g001", "stats"=>{"downloads"=>1, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Separation_of_cell_extracts_on_a_sucrose_gradient_by_ultra_centrifugation_/466819", "title"=>"Separation of cell extracts on a sucrose gradient by ultra-centrifugation.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 01:53:39"}
  • {"files"=>["https://ndownloader.figshare.com/files/796577"], "description"=>"<p>The lower panels depict densitometric scans of the western blots. A) Samples treated with SDS sample buffer at room temperature were SDS-PAGE separated and immunoblotted for DRα (monoclonal antibody TAL-1B5). The upper panel shows SDS resistant pDRαβ bands (ca. 55 kDa) in fractions 6 to 11. The Scan represents both the pDRαβ and the DRα bands. B) Immunoprecipitation (IP) of DRαβ with monoclonal antibody 16.23, and western blotting for DRα (upper panel). The monoclonal antibody 16.23 selectively immunoisolates DR from fraction 6 to 10 and shows no reactivity to DR in fractions 11 to 17. C) Immunoprecipitation from sucrose gradient fractions 6 to 17 with DR monoclonal antibody ISCR3. SDS-PAGE separation and western blotting for DRα (monoclonal antibody TAL-1B5) results in separation of two DR populations, which is visualized as two separate curves in the density scan.</p>", "links"=>[], "tags"=>["raji", "extracts", "sucrose", "gradient"], "article_id"=>466946, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g002", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_SDS_PAGE_separation_of_Raji_cell_extracts_from_sucrose_gradient_fractions_/466946", "title"=>"SDS-PAGE separation of Raji cell extracts from sucrose gradient fractions.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 01:55:46"}
  • {"files"=>["https://ndownloader.figshare.com/files/398443", "https://ndownloader.figshare.com/files/398479", "https://ndownloader.figshare.com/files/398508"], "description"=>"<div><h3>Background</h3><p>The HLA gene complex encodes three class II isotypes, DR, DQ, and DP. HLA class II molecules are peptide receptors that present antigens for recognition by T lymphocytes. In antigen presenting cells, the assembly of matched α and β subunits to heterodimers is chaperoned by invariant chain (Ii). Ii forms a homotrimer with three binding sites for class II heterodimers. The current model of class II and Ii structure states that three αβ heterodimers bind to an Ii trimer.</p> <h3>Methology/Principal Findings</h3><p>We have now analyzed the composition and size of the complexes of class II and Ii using epitope tagged class II subunits and density gradient experiments. We show here that class II-Ii oligomers consist of one class II heterodimer associated with one Ii trimer, such that the DR, DQ and DP isotypes are contained within separate complexes with Ii.</p> <h3>Conclusion/Significance</h3><p>We propose a structural model of the class II-Ii oligomer and speculate that the pentameric class II-Ii complex is bent towards the cell membrane, inhibiting the binding of additional class II heterodimers to Ii.</p> </div>", "links"=>[], "tags"=>["stoichiometry", "hla", "ii-invariant", "oligomers"], "article_id"=>138711, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0017257.s001", "https://dx.doi.org/10.1371/journal.pone.0017257.s002", "https://dx.doi.org/10.1371/journal.pone.0017257.s003"], "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Stoichiometry_of_HLA_Class_II_Invariant_Chain_Oligomers/138711", "title"=>"Stoichiometry of HLA Class II-Invariant Chain Oligomers", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2011-02-22 02:25:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/796992"], "description"=>"<p>A) MelJuSo cells were transiently transfected with cDNAs encoding V5-tagged DPα or DPβ encoding cDNAs. Cells lysed in 1% NP40 were immunoprecipitated for DRαβ (monoclonal antibody ISCR3) (lanes 1,2,3,5,6 and 7). Lanes 4 and 8 contain cell lysates. The SDS-separated polypeptides were immunoblotted for DRα (lanes 1 and 5), for Ii (lanes 2 and 6), and for DPαV5 (lanes 3 and 4), or for DPβV5 chains (lanes 7 and 8). The position of DRα, Ii and Ab-epitope tagged DPαV5 or DPβV5 bands is indicated. H and L chain bands derived from the Ab used for immunoprecipitation are labeled on the left. B) Immunoprecipitates of the V5-tagged DPβ (lanes 1 and 4) and of DRαβ (lanes 2 and 5) chains were separated by SDS-PAGE and either blotted for DPβV5 (lanes 1 to 3) or for DRα (lanes 4 and 5). To monitor expression of DPβV5, lane 3 contains cell lysate. The position of DPβV5, of DRα and of H and L Ig chains are indicated. C) MelJuSo cells were transfected with V5-tagged DQα cDNA. Cell lysate was immunoprecipitated for the V5-tagged DQα chains. Immunoprecipitates (lanes 4 to 6) and cell lysates (lanes 1 to 3) were digested with EndoH (E<sub>H</sub>), or with PNGase F (P<sub>F</sub>) or left untreated (Ø), separated by SDS-PAGE and western blotted as indicated. In lanes 7 to 9 MelJuSo cell lysates untreated and glycosidase treated were western blotted for DRα.</p>", "links"=>[], "tags"=>["co-isolation", "dp", "dq", "isotypes"], "article_id"=>467361, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g005", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Inspection_for_co_isolation_of_DP_and_DQ_isotypes_with_DR_/467361", "title"=>"Inspection for co-isolation of DP and DQ isotypes with DR.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:02:41"}
  • {"files"=>["https://ndownloader.figshare.com/files/797340"], "description"=>"<p>(left) Molecular model (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0017257#s4\" target=\"_blank\">Methods</a> section for details) for a pentameric MHCII-Ii oligomer consisting of a trimeric Ii (chains are in blue, red and green cartoons) and one bound MHC-II heterodimer (alpha and beta chains in pink and brown cartoon, respectively). The position of the membrane is indicated (bars representing approximately the lipid-solvent boundary). (right) Top view of the model (from the ER) along the axis of the coiled-coil structure.</p>", "links"=>[], "tags"=>["mhcii-ii"], "article_id"=>467711, "categories"=>["Immunology"], "users"=>["Norbert Koch", "Martin Zacharias", "Angelika König", "Sebastian Temme", "Jürgen Neumann", "Sebastian Springer"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0017257.g009", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Modeling_of_MHCII_Ii_structure_/467711", "title"=>"Modeling of MHCII-Ii structure.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-02-22 02:08:31"}

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  • {"unique-ip"=>"4", "full-text"=>"2", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2014", "month"=>"8"}
  • {"unique-ip"=>"4", "full-text"=>"0", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2014", "month"=>"9"}
  • {"unique-ip"=>"6", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2014", "month"=>"10"}
  • {"unique-ip"=>"5", "full-text"=>"1", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"2"}
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  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"3", "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"=>"4", "full-text"=>"3", "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"=>"2", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "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"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"4", "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"=>"10"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"3", "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"=>"4", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"12"}
  • {"unique-ip"=>"2", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"2"}
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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"=>"2017", "month"=>"5"}
  • {"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"=>"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"=>"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"=>"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"=>"2017", "month"=>"9"}
  • {"unique-ip"=>"3", "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"=>"10"}
  • {"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"=>"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"=>"2017", "month"=>"12"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "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"=>"2018", "month"=>"2"}
  • {"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"=>"2018", "month"=>"3"}
  • {"unique-ip"=>"5", "full-text"=>"6", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"1"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"18", "full-text"=>"15", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"6", "cited-by"=>"0", "year"=>"2018", "month"=>"5"}
  • {"unique-ip"=>"2", "full-text"=>"0", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"6"}
  • {"unique-ip"=>"8", "full-text"=>"5", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"0", "year"=>"2018", "month"=>"7"}
  • {"unique-ip"=>"7", "full-text"=>"4", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"3", "cited-by"=>"0", "year"=>"2018", "month"=>"8"}
  • {"unique-ip"=>"5", "full-text"=>"5", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"10"}
  • {"unique-ip"=>"1", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2018", "month"=>"9"}
  • {"unique-ip"=>"5", "full-text"=>"4", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"12"}
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  • {"unique-ip"=>"9", "full-text"=>"14", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"2"}
  • {"unique-ip"=>"4", "full-text"=>"2", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"42", "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"=>"7", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "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"=>"12", "full-text"=>"9", "pdf"=>"5", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"9"}
  • {"unique-ip"=>"3", "full-text"=>"3", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"10"}
  • {"unique-ip"=>"8", "full-text"=>"4", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"4", "cited-by"=>"0", "year"=>"2019", "month"=>"12"}
  • {"unique-ip"=>"3", "full-text"=>"1", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"4", "cited-by"=>"0", "year"=>"2020", "month"=>"2"}
  • {"unique-ip"=>"4", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"6", "cited-by"=>"0", "year"=>"2020", "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"=>"2020", "month"=>"4"}
  • {"unique-ip"=>"3", "full-text"=>"1", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "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"=>"2020", "month"=>"6"}
  • {"unique-ip"=>"2", "full-text"=>"5", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"7"}
  • {"unique-ip"=>"2", "full-text"=>"2", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"8"}

Relative Metric

{"start_date"=>"2011-01-01T00:00:00Z", "end_date"=>"2011-12-31T00:00:00Z", "subject_areas"=>[{"subject_area"=>"/Biology and life sciences", "average_usage"=>[304, 568, 702, 818, 927, 1027, 1118, 1206, 1285, 1357, 1427, 1500, 1564, 1636, 1705, 1773, 1840, 1909, 1974, 2039, 2106, 2170, 2234, 2296, 2358, 2423, 2484, 2546, 2606, 2673, 2734, 2795, 2857, 2921, 2984, 3046, 3100]}, {"subject_area"=>"/Biology and life sciences/Anatomy and physiology", "average_usage"=>[298, 559, 690, 805, 916, 1018, 1115, 1201, 1283, 1356, 1425, 1498, 1565, 1635, 1703, 1767, 1835, 1902, 1962, 2027, 2089, 2150, 2210, 2265]}, {"subject_area"=>"/Biology and life sciences/Biochemistry", "average_usage"=>[290, 559, 698, 813, 927, 1028, 1121, 1206, 1289, 1360, 1433, 1501, 1569, 1637, 1707, 1774, 1841, 1911, 1974, 2038, 2103, 2161, 2219, 2280, 2345, 2406, 2466, 2529, 2592, 2654, 2713, 2772, 2835, 2893, 2954, 3020, 3071]}, {"subject_area"=>"/Engineering and technology", "average_usage"=>[312, 536, 658, 770, 877, 959, 1047, 1128, 1203, 1267, 1324, 1404, 1481, 1548, 1609, 1671, 1739, 1802, 1854, 1915, 1974, 2027, 2101, 2161, 2218, 2280, 2340, 2406, 2457, 2507, 2565, 2631, 2682, 2749, 2796, 2844, 2892]}, {"subject_area"=>"/Engineering and technology/Signal processing", "average_usage"=>[324, 555, 686, 779, 884, 954, 1015, 1088, 1148, 1205, 1279, 1317, 1377, 1433, 1490, 1539, 1605, 1662, 1716, 1758, 1824, 1885, 1952, 2004, 2055, 2112, 2178, 2232, 2280, 2305, 2348, 2385, 2442, 2486, 2554, 2628, 2641]}, {"subject_area"=>"/Medicine and health sciences/Anatomy and physiology", "average_usage"=>[296, 554, 686, 802, 910, 1014, 1104, 1195, 1277, 1349, 1419, 1483, 1551, 1620, 1690, 1757, 1820, 1888, 1957, 2017, 2081, 2140, 2202, 2259]}, {"subject_area"=>"/Medicine and health sciences/Immunology", "average_usage"=>[299, 582, 725, 844, 971, 1076, 1167, 1253, 1328, 1403, 1480, 1551, 1620, 1691, 1756, 1821, 1893, 1962, 2032, 2110, 2178, 2236, 2300, 2358, 2419, 2465, 2529, 2587, 2643, 2700, 2756, 2817, 2869, 2932, 2986, 3046, 3101]}, {"subject_area"=>"/Physical sciences/Materials science", "average_usage"=>[284, 516, 629, 726, 835, 925, 1006, 1080, 1148, 1224, 1288, 1355, 1434, 1501, 1563, 1633, 1716, 1767, 1833, 1890, 1949, 2016, 2072, 2132, 2181, 2232, 2293, 2341, 2413, 2480, 2546, 2615, 2678, 2736, 2796, 2869, 2929]}]}
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Net::HTTPTooManyRequests

Source
Scopus
Time
2019-08-27 19:05:11 UTC
Target URL
https://api.elsevier.com/content/search/index:SCOPUS?query=DOI(10.1371%2Fjournal.pone.0017257)
Trace

/app/models/concerns/networkable.rb:21:in `get_result'
/app/models/source.rb:165:in `get_data'
/app/models/retrieval_status.rb:47:in `perform_get_data'
/app/jobs/source_job.rb:52:in `block (2 levels) in perform'
/app/jobs/source_job.rb:51:in `block in perform'
/app/jobs/source_job.rb:35:in `each'
/app/jobs/source_job.rb:35:in `perform'