Human SHBG mRNA Translation Is Modulated by Alternative 5′-Non-Coding Exons 1A and 1B
Publication Date
November 04, 2010
Journal
PLOS ONE
Authors
Tomàs Pinós, Anna Barbosa Desongles, Antoni Hurtado, Albert Santamaria Martínez, et al
Volume
5
Issue
11
Pages
e13844
DOI
https://dx.plos.org/10.1371/journal.pone.0013844
Publisher URL
http://journals.plos.org/plosone/article?id=10.1371%2Fjournal.pone.0013844
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/21079794
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2973947
Europe PMC
http://europepmc.org/abstract/MED/21079794
Web of Science
000283838600018
Scopus
78149481755
Mendeley
http://www.mendeley.com/research/human-shbg-mrna-translation-modulated-alternative-5noncoding-exons-1a-1b
Events
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Mendeley | Further Information

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Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/820295"], "description"=>"<p>A) RT-PCR analysis of SHBG expression in human prostate-derived cell lines, using primers that amplify a region common to all SHBG mRNA isoforms (Ex2-Ex5). B) RT-PCR analysis (Ex1A-Ex8) of exon 1A transcripts in LNCaP cells. The different bands observed correspond to: α (full length transcript), β (skipping of exon 7) and γ (skipping of exons 6 and 7). C) As previously reported <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0013844#pone.0013844-Pinos1\" target=\"_blank\">[2]</a>, full-length TU-1B transcripts are detected by RT-PCR analysis (Ex1B-Ex8) in human prostate tissue and LNCaP cells. Negative controls (Ctrl) are performed with water instead of cDNA.</p>", "links"=>[], "tags"=>["prostate"], "article_id"=>490659, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g004", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_SHBG_expression_in_human_prostate_cell_lines_/490659", "title"=>"SHBG expression in human prostate cell lines.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:10:59"}
  • {"files"=>["https://ndownloader.figshare.com/files/820103"], "description"=>"<p>M-fold prediction of the 5′UTR mRNA secondary structure of TU-1 (A) and TU-1A (B). Predicted thermodynamic stability is shown, and the first SHBG codon is marked with a grey box. Junction between exons 1A and 2 is indicated with a solid black line in B. The uAUG is shown in a blue box in B. The predicted major hairpin is also indicated (HP).</p>", "links"=>[], "tags"=>["sequences", "tu-1"], "article_id"=>490468, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g002", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Predicted_secondary_structure_for_5_8242_UTR_sequences_of_TU_1_and_TU_1A_/490468", "title"=>"Predicted secondary structure for 5′UTR sequences of TU-1 and TU-1A.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:07:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/408871", "https://ndownloader.figshare.com/files/408910", "https://ndownloader.figshare.com/files/408946"], "description"=>"<div><h3>Background</h3><p>The human sex hormone-binding globulin (SHBG) gene comprises at least 6 different transcription units (TU-1, -1A, -1B, -1C, -1D and -1E), and is regulated by no less than 6 different promoters. The best characterized are TU-1 and TU-1A: TU-1 is responsible for producing plasma SHBG, while TU-1A is transcribed and translated in the testis. Transcription of the recently described TU-1B, -1C, and -1D has been demonstrated in human prostate tissue and prostate cancer cell lines, as well as in other human cell lines such as HeLa, HepG2, HeK 293, CW 9019 and imr 32. However, there are no reported data demonstrating their translation. In the present study, we aimed to determine whether TU-1A and TU-1B are indeed translated in the human prostate and whether 5′ UTR exons 1A and 1B differently regulate SHBG translation.</p><h3>Results</h3><p>Cis-regulatory elements that could potentially regulate translation were identified within the 5′UTRs of SHBG TU-1A and TU–1B. Although full-length SHBG TU-1A and TU-1B mRNAs were present in prostate cancer cell lines, the endogenous SHBG protein was not detected by western blot in any of them. LNCaP prostate cancer cells transfected with several SHBG constructs containing exons 2 to 8 but lacking the 5′UTR sequence did show SHBG translation, whereas inclusion of the 5′UTR sequences of either exon 1A or 1B caused a dramatic decrease in SHBG protein levels. The molecular weight of SHBG did not vary between cells transfected with constructs with or without the 5′UTR sequence, thus confirming that the first in-frame ATG of exon 2 is the translation start site of TU-1A and TU-1B.</p><h3>Conclusions</h3><p>The use of alternative SHBG first exons 1A and 1B differentially inhibits translation from the ATG situated in exon 2, which codes for methionine 30 of transcripts that begin with the exon 1 sequence.</p></div>", "links"=>[], "tags"=>["shbg", "mrna", "modulated", "exons", "1a", "1b"], "article_id"=>140773, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0013844.s001", "https://dx.doi.org/10.1371/journal.pone.0013844.s002", "https://dx.doi.org/10.1371/journal.pone.0013844.s003"], "stats"=>{"downloads"=>2, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Human_SHBG_mRNA_Translation_Is_Modulated_by_Alternative_5_Non_Coding_Exons_1A_and_1B/140773", "title"=>"Human SHBG mRNA Translation Is Modulated by Alternative 5′-Non-Coding Exons 1A and 1B", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2010-11-04 00:12:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/819947"], "description"=>"<p>A) The upper panel contains a schematic representation of the SHBG gene. Coding exons are shown in black boxes and noncoding alternative first exons are in gray boxes, with their respective sizes (in bp) indicated above. Below are depicted the exon organization of mature mRNAs of TU-1A and 1B and the corresponding ORF organization. uAUGs (AUG1 and 2) are written in gray while SHBG AUG is written in black (AUG3). uORFs are indicated in gray boxes while SHBG ORF is shown in a black box. AUG1 generates an uORF of 2 codons while AUG2 generates an uORF of 11 codons. B). Exon 2 scheme, showing AUG2 and 3. AUG2 starts a uORF whose stop codon overlaps the SHBG translation start codon (AUG3).</p>", "links"=>[], "tags"=>["uorfs", "tu-1a"], "article_id"=>490311, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g001", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Identification_of_uORFs_in_TU_1A_and_TU_1B_/490311", "title"=>"Identification of uORFs in TU-1A and TU-1B.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:05:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/820373"], "description"=>"<p>A–B) Two bands sized 52- and 48-kDa were detected in human prostate tissue using the SHBG 11F11 antibody (A) and were identical in size to those detected in human plasma and human testis (B). C) SHBG constructs used to transfect LNCaP cells. In the Flag-Ex2-Ex8 construct, SHBG ATG was mutated to ATA to avoid disturbing the Flag translation start site. The nucleotides involved in the SHBG translation start site are shown in all the constructs. D−E) Western blot analysis of the SHBG protein using the 11F11 antibody in transfected and non-transfected human prostate-derived cell lines (D), and in the supernatant (E). In (E), plasma and the hepatocarcinoma cell line (HepG2) were used as positive controls for secreted SHBG. F) Western blot analysis of LNCaP cells transfected or not with Flag-SHBG and pDsRed-SHBG constructs.</p>", "links"=>[], "tags"=>["blot", "shbg"], "article_id"=>490736, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g005", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Western_blot_analysis_of_SHBG_protein_in_human_prostate_/490736", "title"=>"Western blot analysis of SHBG protein in human prostate.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:12:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/820757"], "description"=>"<p>List of primers used for RT-PCR.</p>", "links"=>[], "tags"=>["primers"], "article_id"=>491121, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.t002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_List_of_primers_used_for_RT_PCR_/491121", "title"=>"List of primers used for RT-PCR.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-11-04 00:18:41"}
  • {"files"=>["https://ndownloader.figshare.com/files/820506"], "description"=>"<p>A) Western blot analysis of SHBG translation in LNCaP cells transfected with different SHBG-pDsRed constructs, with and without different 5′UTRs (exon 1A or exon 1B). Human plasma and Flag-SHBG constructs were used as positive controls of SHBG protein detection, while LNCaP cells transfected with pDsRed empty vector and non-transfected cells were used as negative controls. β-actin was used to normalize the quantity of protein loaded on the acrylamide gel. RT-PCR analysis using primers that recognize a region common to all SHBG isoforms (Ex2-Ex8) was performed to normalize the transfection levels with the different SHBG constructs. S18 primers were used to normalize cDNA levels loaded into the PCR reaction. B) Relative protein levels (RPL) and relative protein/mRNA levels (RPML) are shown. RPL was calculated as a ratio of the β-actin level in the sample. RPML was obtained by dividing RPL by the quantified intensity of mRNA SHBG bands. C) Western blot analysis of SHBG translation in LNCaP cells transfected with pcDNA 3-SHBG constructs with and without the 5′UTR (exon 1A). D) RPL and RPML were calculated as above.</p>", "links"=>[], "tags"=>["differentially", "modulated", "exons", "1a"], "article_id"=>490868, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g006", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_SHBG_translation_is_differentially_modulated_by_alternative_exons_1A_and_1B_/490868", "title"=>"SHBG translation is differentially modulated by alternative exons 1A and 1B.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:14:28"}
  • {"files"=>["https://ndownloader.figshare.com/files/820725"], "description"=>"<p>SHBG classification of alternative 5′UTRs.</p>", "links"=>[], "tags"=>["classification"], "article_id"=>491093, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.t001", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_SHBG_classification_of_alternative_5_8242_UTRs_/491093", "title"=>"SHBG classification of alternative 5′UTRs.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-11-04 00:18:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/820644"], "description"=>"<p>A) Western blot analysis of SHBG translation in LNCaP cells transfected with pcDNA 3-SHBG constructs with and without 5′UTR (exon 1A). Three independent experiments were performed. B) RPL and RPML were calculated. C) Western blot analysis of SHBG translation in DU-145 cells transfected with pcDNA 3-SHBG constructs with and without the 5′UTR (exon 1A). Three independent experiments were performed. D) RPL and RPML were calculated.</p>", "links"=>[], "tags"=>["pcdna", "constructs", "lncap", "du-145"], "article_id"=>491003, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g007", "stats"=>{"downloads"=>1, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Transfection_of_pcDNA_3_constructs_in_LNCaP_and_DU_145_cell_lines_/491003", "title"=>"Transfection of pcDNA 3 constructs in LNCaP and DU-145 cell lines.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:16:43"}
  • {"files"=>["https://ndownloader.figshare.com/files/820221"], "description"=>"<p>M-fold prediction of the 5′UTR mRNA secondary structure of TU-1B. Predicted thermodynamic stability, the first SHBG codon, junction between exons 1B and 2, and the uAUG are indicated as in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0013844#pone-0013844-g002\" target=\"_blank\">Figure 2</a>. The presence of hairpins is also indicated (HP1, HP2, HP3, HP4 and HP5).</p>", "links"=>[], "tags"=>["sequences"], "article_id"=>490589, "categories"=>["Molecular Biology", "Medicine", "Genetics"], "users"=>["Tomàs Pinós", "Anna Barbosa-Desongles", "Antoni Hurtado", "Albert Santamaria-Martínez", "Inés de Torres", "Jaume Reventós", "Francina Munell"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0013844.g003", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Predicted_secondary_structure_for_5_8242_UTR_sequences_of_TU_1B_/490589", "title"=>"Predicted secondary structure for 5′UTR sequences of TU-1B.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-11-04 00:09:49"}

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

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