Clusters of Conserved Beta Cell Marker Genes for Assessment of Beta Cell Phenotype
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{"title"=>"Clusters of conserved beta cell marker genes for assessment of beta cell phenotype", "type"=>"journal", "authors"=>[{"first_name"=>"Geert A.", "last_name"=>"Martens", "scopus_author_id"=>"8592742200"}, {"first_name"=>"Lei", "last_name"=>"Jiang", "scopus_author_id"=>"57193925625"}, {"first_name"=>"Karine H.", "last_name"=>"Hellemans", "scopus_author_id"=>"6603516224"}, {"first_name"=>"Geert", "last_name"=>"Stangé", "scopus_author_id"=>"7005420751"}, {"first_name"=>"Harry", "last_name"=>"Heimberg", "scopus_author_id"=>"7003495114"}, {"first_name"=>"Finn C.", "last_name"=>"Nielsen", "scopus_author_id"=>"7201827789"}, {"first_name"=>"Olivier", "last_name"=>"Sand", "scopus_author_id"=>"24081512500"}, {"first_name"=>"Jacques", "last_name"=>"van Helden", "scopus_author_id"=>"7003886643"}, {"first_name"=>"Frans K.", "last_name"=>"Gorus", "scopus_author_id"=>"7005753962"}, {"first_name"=>"Daniel G.", "last_name"=>"Pipeleers", "scopus_author_id"=>"7102181294"}], "year"=>2011, "source"=>"PLoS ONE", "identifiers"=>{"scopus"=>"2-s2.0-80052407683", "sgr"=>"80052407683", "issn"=>"19326203", "pmid"=>"21912665", "doi"=>"10.1371/journal.pone.0024134", "pui"=>"362489472"}, "id"=>"85ff05cb-8dcd-3a9b-a674-f76baebe5e54", "abstract"=>"BACKGROUND AND METHODOLOGY: The aim of this study was to establish a gene expression blueprint of pancreatic beta cells conserved from rodents to humans and to evaluate its applicability to assess shifts in the beta cell differentiated state. Genome-wide mRNA expression profiles of isolated beta cells were compared to those of a large panel of other tissue and cell types, and transcripts with beta cell-abundant and -selective expression were identified. Iteration of this analysis in mouse, rat and human tissues generated a panel of conserved beta cell biomarkers. This panel was then used to compare isolated versus laser capture microdissected beta cells, monitor adaptations of the beta cell phenotype to fasting, and retrieve possible conserved transcriptional regulators.\\n\\nPRINCIPAL FINDINGS: A panel of 332 conserved beta cell biomarker genes was found to discriminate both isolated and laser capture microdissected beta cells from all other examined cell types. Of all conserved beta cell-markers, 15% were strongly beta cell-selective and functionally associated to hormone processing, 15% were shared with neuronal cells and associated to regulated synaptic vesicle transport and 30% with immune plus gut mucosal tissues reflecting active protein synthesis. Fasting specifically down-regulated the latter cluster, but preserved the neuronal and strongly beta cell-selective traits, indicating preserved differentiated state. Analysis of consensus binding site enrichment indicated major roles of CREB/ATF and various nutrient- or redox-regulated transcription factors in maintenance of differentiated beta cell phenotype.\\n\\nCONCLUSIONS: Conserved beta cell marker genes contain major gene clusters defined by their beta cell selectivity or by their additional abundance in either neural cells or in immune plus gut mucosal cells. This panel can be used as a template to identify changes in the differentiated state of beta cells.", "link"=>"http://www.mendeley.com/research/clusters-conserved-beta-cell-marker-genes-assessment-beta-cell-phenotype", "reader_count"=>61, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>3, "Student > Doctoral Student"=>4, "Researcher"=>18, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>5, "Student > Master"=>4, "Other"=>4, "Student > Bachelor"=>4, "Lecturer > Senior Lecturer"=>1, "Professor"=>3}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>3, "Student > Doctoral Student"=>4, "Researcher"=>18, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>5, "Student > Master"=>4, "Other"=>4, "Student > Bachelor"=>4, "Lecturer > Senior Lecturer"=>1, "Professor"=>3}, "reader_count_by_subject_area"=>{"Unspecified"=>3, "Biochemistry, Genetics and Molecular Biology"=>10, "Agricultural and Biological Sciences"=>37, "Medicine and Dentistry"=>10, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>10}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>37}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>10}, "Unspecified"=>{"Unspecified"=>3}}, "reader_count_by_country"=>{"Czech Republic"=>1, "Belgium"=>1, "Israel"=>1, "Jordan"=>1, "France"=>1, "Switzerland"=>1, "Germany"=>1, "Spain"=>2}, "group_count"=>4}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/739079"], "description"=>"<p>Beta cell preparations were compared to the indicated tissues in mouse, rat and human and genes with conserved beta cell-abundant expression selected (Venn diagram intersect). The rat data set was used to discard 46 islet endocrine non-beta cell (mainly alpha cell) markers; the human data set enabled curation of 17 pancreatic exocrine (acinar and duct) markers. Final set of conserved beta cell marker genes consists of 332 genes, represented by 419 (rat), 499 (mouse) or 503 (human) oligonucleotide probe sets.</p>", "links"=>[], "tags"=>["algorithm", "conserved", "beta"], "article_id"=>409440, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g001", "stats"=>{"downloads"=>0, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Selection_algorithm_of_conserved_beta_cell_marker_genes_/409440", "title"=>"Selection algorithm of conserved beta cell marker genes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 02:37:20"}
  • {"files"=>["https://ndownloader.figshare.com/files/739207"], "description"=>"<p>The 503 human probe sets corresponding to the 332 conserved beta cell marker genes are visualized by hierarchical clustering in the Human Genome Atlas data set <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone.0024134-Su1\" target=\"_blank\">[3]</a>. At least 15% of beta cell marker genes show near absolute beta cell selectivity (beta cell selective <b>cluster B</b>). Beta cells also share an equally extensive cluster (15%) with various brain regions (neuroendocrine <b>cluster A</b>), and an even larger cluster (30%) with immune and gut mucosal cells (gut/immunological <b>cluster C</b>). Statistical enrichment analysis indicates that these clusters accentuate different functions proper to the specialized beta cell phenotype. Blue to red correspond low to high relative expression, as shown in legend (bottom). Relevant genes of clusters A, B and C are shown in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone-0024134-g003\" target=\"_blank\">Fig. 3</a>.</p>", "links"=>[], "tags"=>["clusters", "beta", "neuronal", "cells"], "article_id"=>409566, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g002", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Shared_gene_clusters_between_beta_cells_neuronal_cells_and_immune_cells_/409566", "title"=>"Shared gene clusters between beta cells, neuronal cells and immune cells.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 02:39:26"}
  • {"files"=>["https://ndownloader.figshare.com/files/740168"], "description"=>"<p>mRNA expression levels are expressed as geometric mean (GEO) ± SD (n = 3) of indicated markers normalized to 4 reference genes (<i>BACT, UBC, PPIA, PSMC5</i>). Marker genes are sorted by decreasing order of their mRNA abundance in beta cells. Duod., duodenum; WAT, white adipose tissue (abdominal); WBC, white blood cells.</p>", "links"=>[], "tags"=>["beta", "genes"], "article_id"=>410533, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.t001", "stats"=>{"downloads"=>1, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_qPCR_analysis_of_beta_cell_marker_genes_in_rat_tissues_/410533", "title"=>"qPCR analysis of beta cell marker genes in rat tissues.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2011-09-02 00:08:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/740130"], "description"=>"<p>Data represent relative molar amounts of proteins coded by beta cell marker genes that could be detected in label-free alternate-scanning LC-MS proteomics of unfractionated cells and tissues (mean of triplicate injections of 3 biological replicates, total CV%<20% <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone.0024134-Martens1\" target=\"_blank\">[9]</a>). For comparison avFC and rank of mRNA transcript (array) are compared to protein abundance (fold above lower limit of quantification, LLQ) and rank (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#s4\" target=\"_blank\">Methods</a> and reference <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone.0024134-Martens1\" target=\"_blank\">[9]</a> for details).</p>", "links"=>[], "tags"=>["beta", "genes", "alpha", "quantitative"], "article_id"=>410490, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.t002", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Conserved_beta_cell_marker_genes_identified_in_rat_beta_cells_alpha_cells_liver_and_brain_tissue_using_quantitative_proteomics_/410490", "title"=>"Conserved beta cell marker genes identified in rat beta cells, alpha cells, liver and brain tissue using quantitative proteomics.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2011-09-02 00:08:10"}
  • {"files"=>["https://ndownloader.figshare.com/files/739311"], "description"=>"<p>Average fold change (AvFC) indicates the relative abundance of a given mRNA in the beta cell as compared to the other tested tissues in that species, while lower rank value reflects higher beta cell selectivity (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#s4\" target=\"_blank\">methods</a>). From each tissue-tropic cluster (A, B, C from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone-0024134-g002\" target=\"_blank\">Fig. 2</a>), beta cell marker genes with, calculated for the 3 species, an averaged avFC score ≥4, are shown. Relative beta cell abundance (and selectivity) decreases from cluster B over cluster A to cluster C, and consequently this is reflected in number of retained exemplary genes.</p>", "links"=>[], "tags"=>["beta", "genes", "belonging", "neuroendocrine", "selective", "beta-immune"], "article_id"=>409667, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g003", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Examples_of_beta_cell_marker_genes_belonging_to_neuroendocrine_cluster_A_beta_cell_selective_cluster_B_and_beta_immune_cluster_C_/409667", "title"=>"Examples of beta cell marker genes belonging to neuroendocrine cluster A, beta cell selective cluster B and beta-immune cluster C.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 02:41:07"}
  • {"files"=>["https://ndownloader.figshare.com/files/739921"], "description"=>"<p>Panels A and B respectively show abundant expression of PPP1R1A protein in insulin+ cells and absence in glucagon+ cells in islets from 10 w old rats. Beta cell-selective expression of PTPRN and HADH proteins in human beta cells is illustrated in panels C and D respectively. All images are representative of at least 3 different organs with at least 3 sections analyzed per pancreas.</p>", "links"=>[], "tags"=>["beta", "cell-selective", "ptprn", "hadh"], "article_id"=>410286, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g007", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_In_situ_analysis_of_beta_cell_selective_expression_of_PPP1R1A_PTPRN_and_HADH_proteins_/410286", "title"=>"<i>In situ</i> analysis of beta cell-selective expression of PPP1R1A, PTPRN and HADH proteins.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 00:04:46"}
  • {"files"=>["https://ndownloader.figshare.com/files/739748"], "description"=>"<p>Using two complementary search algorithms (Matrix Scan and DiRE), upstream promoter regions were scanned for statistical enrichment of TRANSFAC-annotated transcription factor binding consensus sites. Venn diagrams indicate number of identified matrices by each program, and with 30 commonly identified matrices indicated in gray field. See <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone.0024134.s007\" target=\"_blank\">Tables S3</a> and <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone.0024134.s008\" target=\"_blank\">S4</a>.</p>", "links"=>[], "tags"=>["binding", "sites", "overrepresented", "conserved", "beta"], "article_id"=>410108, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g006", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Transcription_factor_binding_sites_overrepresented_in_the_conserved_beta_cell_marker_gene_set_/410108", "title"=>"Transcription factor binding sites overrepresented in the conserved beta cell marker gene set.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 00:01:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/373039", "https://ndownloader.figshare.com/files/373089", "https://ndownloader.figshare.com/files/373122", "https://ndownloader.figshare.com/files/373154", "https://ndownloader.figshare.com/files/373194", "https://ndownloader.figshare.com/files/373369", "https://ndownloader.figshare.com/files/373572", "https://ndownloader.figshare.com/files/373618"], "description"=>"<div><h3>Background and Methodology</h3><p>The aim of this study was to establish a gene expression blueprint of pancreatic beta cells conserved from rodents to humans and to evaluate its applicability to assess shifts in the beta cell differentiated state. Genome-wide mRNA expression profiles of isolated beta cells were compared to those of a large panel of other tissue and cell types, and transcripts with beta cell-abundant and -selective expression were identified. Iteration of this analysis in mouse, rat and human tissues generated a panel of conserved beta cell biomarkers. This panel was then used to compare isolated versus laser capture microdissected beta cells, monitor adaptations of the beta cell phenotype to fasting, and retrieve possible conserved transcriptional regulators.</p> <h3>Principal Findings</h3><p>A panel of 332 conserved beta cell biomarker genes was found to discriminate both isolated and laser capture microdissected beta cells from all other examined cell types. Of all conserved beta cell-markers, 15% were strongly beta cell-selective and functionally associated to hormone processing, 15% were shared with neuronal cells and associated to regulated synaptic vesicle transport and 30% with immune plus gut mucosal tissues reflecting active protein synthesis. Fasting specifically down-regulated the latter cluster, but preserved the neuronal and strongly beta cell-selective traits, indicating preserved differentiated state. Analysis of consensus binding site enrichment indicated major roles of CREB/ATF and various nutrient- or redox-regulated transcription factors in maintenance of differentiated beta cell phenotype.</p> <h3>Conclusions</h3><p>Conserved beta cell marker genes contain major gene clusters defined by their beta cell selectivity or by their additional abundance in either neural cells or in immune plus gut mucosal cells. This panel can be used as a template to identify changes in the differentiated state of beta cells.</p> </div>", "links"=>[], "tags"=>["clusters", "conserved", "beta", "genes", "phenotype"], "article_id"=>133674, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024134.s001", "https://dx.doi.org/10.1371/journal.pone.0024134.s002", "https://dx.doi.org/10.1371/journal.pone.0024134.s003", "https://dx.doi.org/10.1371/journal.pone.0024134.s004", "https://dx.doi.org/10.1371/journal.pone.0024134.s005", "https://dx.doi.org/10.1371/journal.pone.0024134.s006", "https://dx.doi.org/10.1371/journal.pone.0024134.s007", "https://dx.doi.org/10.1371/journal.pone.0024134.s008"], "stats"=>{"downloads"=>4, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Clusters_of_Conserved_Beta_Cell_Marker_Genes_for_Assessment_of_Beta_Cell_Phenotype/133674", "title"=>"Clusters of Conserved Beta Cell Marker Genes for Assessment of Beta Cell Phenotype", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2011-09-02 01:01:14"}
  • {"files"=>["https://ndownloader.figshare.com/files/739467"], "description"=>"<p>Panel A shows mRNA expression signal on HG133A chip of 13 established pancreatic exocrine (acinar) genes, in freshly isolated human islets (islets, red bars, n = 3), laser capture microdissected beta cells (LCM, green bars, n = 3) and 2–3 w cultured beta cells that were subsequently FACS-enriched (beta cells, blue bars, n = 3). * p<0.05 versus LCM. Panel B shows relative mRNA levels established beta cell-specific genes in cultured beta (blue), fresh islets (red) and LCM-beta cells (green). Table shows fold change (LCB, lower confidence bound) and P values of versus LCM-beta.</p>", "links"=>[], "tags"=>["beta", "genes", "microdissected"], "article_id"=>409829, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g004", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Conserved_beta_cell_marker_genes_in_isolated_versus_laser_capture_microdissected_human_beta_cells_/409829", "title"=>"Conserved beta cell marker genes in isolated versus laser capture microdissected human beta cells.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 02:43:49"}
  • {"files"=>["https://ndownloader.figshare.com/files/739561"], "description"=>"<p>Heat map in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024134#pone-0024134-g005\" target=\"_blank\">Fig. 5A</a> shows the complete set of species-conserved beta cell markers, visualized in FACS-purified beta cells freshly isolated from control (FED) and 24-h fasted (FAST) rats; one third is reproducibly ≥1.25-times down-regulated (blue window) most of which statistically significantly (21% of all beta markers, p<0.05, n = 3). Table in panel B shows the 42 beta markers that are at least 1.5-fold down-regulated (p<0.05) by fasting: 50% of the fasting-suppressed beta cell marker genes belong to cluster C, also functionally enriched in pathways of protein folding and processing through ER/Golgi.</p>", "links"=>[], "tags"=>["beta", "cells", "fasting", "down-regulates", "genes", "er"], "article_id"=>409925, "categories"=>["Chemistry", "Biological Sciences", "Genetics", "Developmental Biology", "Evolutionary Biology"], "users"=>["Geert A. Martens", "Lei Jiang", "Karine H. Hellemans", "Geert Stangé", "Harry Heimberg", "Finn C. Nielsen", "Olivier Sand", "Jacques van Helden", "Frans K. Gorus", "Daniel G. Pipeleers"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0024134.g005", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_In_rat_beta_cells_fasting_down_regulates_cluster_C_genes_with_role_in_ER_function_and_protein_processing_/409925", "title"=>"In rat beta cells fasting down-regulates cluster C genes with role in ER function and protein processing.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-02 02:45:25"}

PMC Usage Stats | Further Information

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Relative Metric

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