A Genome-Wide Association Study in Chronic Obstructive Pulmonary Disease (COPD): Identification of Two Major Susceptibility Loci
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{"title"=>"A Genome-Wide association study in chronic obstructive pulmonary disease (COPD): Identification of two major susceptibility loci", "type"=>"journal", "authors"=>[{"first_name"=>"Sreekumar G.", "last_name"=>"Pillai", "scopus_author_id"=>"35262529200"}, {"first_name"=>"Dongliang", "last_name"=>"Ge", "scopus_author_id"=>"8963757100"}, {"first_name"=>"Guohua", "last_name"=>"Zhu", "scopus_author_id"=>"55451893700"}, {"first_name"=>"Xiangyang", "last_name"=>"Kong", "scopus_author_id"=>"35976173000"}, {"first_name"=>"Kevin V.", "last_name"=>"Shianna", "scopus_author_id"=>"6508389190"}, {"first_name"=>"Anna C.", "last_name"=>"Need", "scopus_author_id"=>"9746096700"}, {"first_name"=>"Sheng", "last_name"=>"Feng", "scopus_author_id"=>"35725490300"}, {"first_name"=>"Craig P.", "last_name"=>"Hersh", "scopus_author_id"=>"6701515147"}, {"first_name"=>"Per", "last_name"=>"Bakke", "scopus_author_id"=>"7005426134"}, {"first_name"=>"Amund", "last_name"=>"Gulsvik", "scopus_author_id"=>"7004578220"}, {"first_name"=>"Andreas", "last_name"=>"Ruppert", "scopus_author_id"=>"6603959519"}, {"first_name"=>"Karin C.Lødrup", "last_name"=>"Carlsen", "scopus_author_id"=>"6603838951"}, {"first_name"=>"Allen", "last_name"=>"Roses", "scopus_author_id"=>"35352401800"}, {"first_name"=>"Wayne", "last_name"=>"Anderson", "scopus_author_id"=>"35433647400"}, {"first_name"=>"Stephen I.", "last_name"=>"Rennard", "scopus_author_id"=>"35396193900"}, {"first_name"=>"David A.", "last_name"=>"Lomas", "scopus_author_id"=>"35391622600"}, {"first_name"=>"Edwin K.", "last_name"=>"Silverman", "scopus_author_id"=>"7201673300"}, {"first_name"=>"David B.", "last_name"=>"Goldstein", "scopus_author_id"=>"7402147672"}], "year"=>2009, "source"=>"PLoS Genetics", "identifiers"=>{"doi"=>"10.1371/journal.pgen.1000421", "sgr"=>"63449101907", "issn"=>"15537390", "pui"=>"354404859", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)", "pmid"=>"19300482", "scopus"=>"2-s2.0-63449101907"}, "id"=>"17473593-c8eb-32f9-b513-fdc9305a72a1", "abstract"=>"There is considerable variability in the susceptibility of smokers to develop chronic obstructive pulmonary disease (COPD). The only known genetic risk factor is severe deficiency of 1-antitrypsin, which is present in 12% of individuals with COPD. We conducted a genome-wide association study (GWAS) in a homogenous case-control cohort from Bergen, Norway (823 COPD cases and 810 smoking controls) and evaluated the top 100 single nucleotide polymorphisms (SNPs) in the family-based International COPD Genetics Network (ICGN; 1891 Caucasian individuals from 606 pedigrees) study. The polymorphisms that showed replication were further evaluated in 389 subjects from the US National Emphysema Treatment Trial (NETT) and 472 controls from the Normative Aging Study (NAS) and then in a fourth cohort of 949 individuals from 127 extended pedigrees from the Boston Early-Onset COPD population. Logistic regression models with adjustments of covariates were used to analyze the case-control populations. Family-based association analyses were conducted for a diagnosis of COPD and lung function in the family populations. Two SNPs at the -nicotinic acetylcholine receptor (CHRNA 3/5) locus were identified in the genome-wide association study. They showed unambiguous replication in the ICGN family-based analysis and in the NETT case-control analysis with combined p-values of 1.481010, (rs8034191) and 5.741010 (rs1051730). Furthermore, these SNPs were significantly associated with lung function in both the ICGN and Boston Early-Onset COPD populations. The C allele of the rs8034191 SNP was estimated to have a population attributable risk for COPD of 12.2%. The association of hedgehog interacting protein (HHIP) locus on chromosome 4 was also consistently replicated, but did not reach genome-wide significance levels. Genome-wide significant association of the HHIP locus with lung function was identified in the Framingham Heart study (Wilk et al., companion article in this issue of PLoS Genetics; The CHRNA 3/5 and the HHIP loci make a significant contribution to the risk of COPD. CHRNA3/5 is the same locus that has been implicated in the risk of lung cancer.", "link"=>"http://www.mendeley.com/research/genomewide-association-study-chronic-obstructive-pulmonary-disease-copd-identification-two-major-sus-10", "reader_count"=>35, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>5, "Researcher"=>8, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>5, "Student > Postgraduate"=>3, "Student > Master"=>5, "Other"=>2, "Student > Bachelor"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>1}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>5, "Researcher"=>8, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>5, "Student > Postgraduate"=>3, "Student > Master"=>5, "Other"=>2, "Student > Bachelor"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>1}, "reader_count_by_subject_area"=>{"Unspecified"=>4, "Environmental Science"=>1, "Nursing and Health Professions"=>1, "Biochemistry, Genetics and Molecular Biology"=>7, "Medicine and Dentistry"=>12, "Agricultural and Biological Sciences"=>8, "Neuroscience"=>1, "Social Sciences"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>12}, "Neuroscience"=>{"Neuroscience"=>1}, "Social Sciences"=>{"Social Sciences"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>8}, "Nursing and Health Professions"=>{"Nursing and Health Professions"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>7}, "Unspecified"=>{"Unspecified"=>4}, "Environmental Science"=>{"Environmental Science"=>1}}, "reader_count_by_country"=>{"Canada"=>2, "Germany"=>1}, "group_count"=>2}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/904238"], "description"=>"a<p>Logistic regression analyses using age, gender, pack years of smoking and current smoking status as co-variates.</p>b<p>International COPD Genetics Network (ICGN) population. PBAT version 3.6 analyses using age, gender, pack-years of smoking, current smoking status and center as co-variates.</p>c<p>National Emphysema Treatment Trail/Normative aging study (NETT/NAS). Logistic regression using age and pack years of smoking (all the subjects are males and ex-smokers, so adjustments for gender and current smoking status were not included).</p>d<p>Fisher's combined probability test was applied to combine the P-values from Bergen Cohort, ICGN cohort and NETT/NAS study.</p><p>P values in bold are above the genome-wide significance level (p<1.01×10<sup>−7</sup>).</p><p>Numbers in Bold are genome-wide significant (p value<1.01×10<sup>−7</sup>).</p>e<p>Risk alleles were different in the ICGN or NETT/NAS population compared to the Bergen discovery cohort.</p>", "links"=>[], "tags"=>["replication", "analyses", "100", "snps"], "article_id"=>574672, "categories"=>["Genetics", "Plant Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000421.t002", "stats"=>{"downloads"=>3, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Results_of_the_replication_association_analyses_of_the_top_100_SNPs_in_the_Norway_ICGN_and_NETT_NAS_populations_/574672", "title"=>"Results of the replication association analyses of the top 100 SNPs in the Norway, ICGN, and NETT/NAS populations.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-03-20 01:17:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/904053"], "description"=>"<p>Annotated sections: Top: Linkage disequilibrium (r<sup>2</sup>) for all HapMap SNPs with rs8034191, showing an LD extension covering CHRNA3/CHRNA5. [red] r<sup>2</sup>≥0.8, [yellow] 0.5≤r2<0.8, [gray] 0.3≤r<sup>2</sup><0.5, [blue] 0.2≤r2<0.3. Middle: Recent selection score. Bottom: genic context. Exons are depicted as blue vertical lines/rectangles, while introns are depicted as blue horizontal lines. Annotations were done using the WGAViewer software <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1000421#pgen.1000421-Ge1\" target=\"_blank\">[31]</a>.</p>", "links"=>[], "tags"=>["genetics and genomics/gene discovery", "respiratory medicine/copd and allied disorders", "respiratory medicine/lung cancer"], "article_id"=>574497, "categories"=>["Genetics", "Plant Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000421.g002", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Region_of_Association_around_CHRNA3_CHRNA5_/574497", "title"=>"Region of Association around CHRNA3/CHRNA5.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-03-20 01:14:57"}
  • {"files"=>["https://ndownloader.figshare.com/files/904298"], "description"=>"<p>Note 1: Heavy smoker defined by pack-years of smoking (grouped by log transformed median = 3.11).</p><p>Note 2: Heavy smoker defined by length of smoking history (grouped by median = 41.5 yr).</p>", "links"=>[], "tags"=>["counts", "allele", "frequencies", "rs8034191", "case-control", "smoking", "bergen"], "article_id"=>574727, "categories"=>["Genetics", "Plant Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000421.t003", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Genotype_counts_and_allele_frequencies_for_rs8034191_by_case_control_status_and_smoking_status_in_the_Bergen_discovery_cohort_/574727", "title"=>"Genotype counts and allele frequencies for rs8034191 by case-control status and smoking status in the Bergen discovery cohort.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-03-20 01:18:47"}
  • {"files"=>["https://ndownloader.figshare.com/files/904170"], "description"=>"a<p>Note: FEV<sub>1</sub>/VC was used for the ICGN Family Data, with VC determined by the higher of FVC and SVC.</p>b<p>Pre-bronchodilator spirometry measurements were used in the NAS.</p>c<p>Post-bronchodilator spirometry measurements available in 118 probands and 789 relatives.</p>", "links"=>[], "tags"=>["replication"], "article_id"=>574609, "categories"=>["Genetics", "Plant Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000421.t001", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Characteristics_of_the_primary_screening_and_replication_populations_/574609", "title"=>"Characteristics of the primary screening and replication populations.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-03-20 01:16:49"}
  • {"files"=>["https://ndownloader.figshare.com/files/903895"], "description"=>"<p>SNP: Single Nucleotide Polymorphism. <sup>a</sup>ICGN: International COPD Genetics Network families. <sup>b</sup>NETT: National Emphysema Treatment Trial. <sup>c</sup>NAS: Normative aging study. <sup>d</sup> One of the SNPs genotyped in NETT/NAS population (rs735243) was not successfully genotyped in the BEOCOPD population.</p>", "links"=>[], "tags"=>["genetics and genomics/gene discovery", "respiratory medicine/copd and allied disorders", "respiratory medicine/lung cancer"], "article_id"=>574329, "categories"=>["Genetics", "Plant Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000421.g001", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Study_design_/574329", "title"=>"Study design.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-03-20 01:12:09"}
  • {"files"=>["https://ndownloader.figshare.com/files/447043", "https://ndownloader.figshare.com/files/447050", "https://ndownloader.figshare.com/files/447062", "https://ndownloader.figshare.com/files/447075", "https://ndownloader.figshare.com/files/447086", "https://ndownloader.figshare.com/files/447093", "https://ndownloader.figshare.com/files/447104"], "description"=>"<div><p>There is considerable variability in the susceptibility of smokers to develop chronic obstructive pulmonary disease (COPD). The only known genetic risk factor is severe deficiency of α<sub>1</sub>-antitrypsin, which is present in 1–2% of individuals with COPD. We conducted a genome-wide association study (GWAS) in a homogenous case-control cohort from Bergen, Norway (823 COPD cases and 810 smoking controls) and evaluated the top 100 single nucleotide polymorphisms (SNPs) in the family-based International COPD Genetics Network (ICGN; 1891 Caucasian individuals from 606 pedigrees) study. The polymorphisms that showed replication were further evaluated in 389 subjects from the US National Emphysema Treatment Trial (NETT) and 472 controls from the Normative Aging Study (NAS) and then in a fourth cohort of 949 individuals from 127 extended pedigrees from the Boston Early-Onset COPD population. Logistic regression models with adjustments of covariates were used to analyze the case-control populations. Family-based association analyses were conducted for a diagnosis of COPD and lung function in the family populations. Two SNPs at the α-nicotinic acetylcholine receptor (CHRNA 3/5) locus were identified in the genome-wide association study. They showed unambiguous replication in the ICGN family-based analysis and in the NETT case-control analysis with combined <em>p</em>-values of 1.48×10<sup>−10</sup>, (rs8034191) and 5.74×10<sup>−10</sup> (rs1051730). Furthermore, these SNPs were significantly associated with lung function in both the ICGN and Boston Early-Onset COPD populations. The C allele of the rs8034191 SNP was estimated to have a population attributable risk for COPD of 12.2%. The association of hedgehog interacting protein (HHIP) locus on chromosome 4 was also consistently replicated, but did not reach genome-wide significance levels. Genome-wide significant association of the HHIP locus with lung function was identified in the Framingham Heart study (Wilk et al., companion article in this issue of <em>PLoS Genetics</em>; doi:10.1371/journal.pgen.1000429). The CHRNA 3/5 and the HHIP loci make a significant contribution to the risk of COPD. CHRNA3/5 is the same locus that has been implicated in the risk of lung cancer.</p></div>", "links"=>[], "tags"=>["genome-wide", "obstructive", "pulmonary", "susceptibility", "loci"], "article_id"=>148119, "categories"=>["Genetics", "Cell Biology", "Cancer"], "users"=>["Sreekumar G. Pillai", "Dongliang Ge", "Guohua Zhu", "Xiangyang Kong", "Kevin V. Shianna", "Anna C. Need", "Sheng Feng", "Craig P. Hersh", "Per Bakke", "Amund Gulsvik", "Andreas Ruppert", "Karin C. Lødrup Carlsen", "Allen Roses", "Wayne Anderson", "Stephen I. Rennard", "David A. Lomas", "Edwin K. Silverman", "David B. Goldstein"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1000421.s001", "https://dx.doi.org/10.1371/journal.pgen.1000421.s002", "https://dx.doi.org/10.1371/journal.pgen.1000421.s003", "https://dx.doi.org/10.1371/journal.pgen.1000421.s004", "https://dx.doi.org/10.1371/journal.pgen.1000421.s005", "https://dx.doi.org/10.1371/journal.pgen.1000421.s006", "https://dx.doi.org/10.1371/journal.pgen.1000421.s007"], "stats"=>{"downloads"=>27, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/A_Genome_Wide_Association_Study_in_Chronic_Obstructive_Pulmonary_Disease_COPD_Identification_of_Two_Major_Susceptibility_Loci/148119", "title"=>"A Genome-Wide Association Study in Chronic Obstructive Pulmonary Disease (COPD): Identification of Two Major Susceptibility Loci", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2009-03-20 02:15:19"}

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  • {"unique-ip"=>"22", "full-text"=>"26", "pdf"=>"4", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"8"}

Relative Metric

{"start_date"=>"2009-01-01T00:00:00Z", "end_date"=>"2009-12-31T00:00:00Z", "subject_areas"=>[{"subject_area"=>"/Biology and life sciences", "average_usage"=>[349, 614, 744, 851, 955, 1059, 1167, 1262, 1352, 1431, 1516, 1593, 1668, 1746, 1817, 1883, 1950, 2017, 2075, 2141, 2198, 2254, 2313, 2368, 2424, 2474, 2534, 2591, 2651, 2710, 2776, 2837, 2892, 2953, 3014, 3072, 3130, 3186, 3251, 3307, 3366, 3427, 3500, 3561, 3627, 3688, 3759, 3821, 3888, 3950, 4010, 4070, 4131, 4189, 4242, 4307, 4370, 4431, 4491, 4549, 4608]}, {"subject_area"=>"/Biology and life sciences/Behavior", "average_usage"=>[407, 682, 829, 937, 1026, 1116, 1212, 1315, 1397, 1487, 1556, 1630, 1701, 1772, 1845, 1902, 2007, 2098, 2155, 2225, 2292, 2350, 2408, 2467, 2496, 2530, 2594, 2656, 2712, 2778, 2860, 2945, 3047, 3099, 3169, 3234, 3311, 3344, 3379, 3422, 3485, 3548, 3594, 3642, 3722, 3789, 3862, 3959, 4067, 4116, 4191, 4265, 4385, 4453, 4535, 4614, 4654, 4719, 4773, 4816, 4878]}, {"subject_area"=>"/Biology and life sciences/Genetics", "average_usage"=>[366, 637, 775, 896, 999, 1118, 1230, 1322, 1416, 1516, 1602, 1685, 1767, 1849, 1914, 1982, 2049, 2120, 2185, 2246, 2303, 2363, 2409, 2473, 2535, 2599, 2655, 2717, 2777, 2842, 2906, 2977, 3021, 3088, 3144, 3199, 3253, 3317, 3379, 3456, 3508, 3567, 3634, 3716, 3781, 3853, 3911, 3974, 4040, 4118, 4170, 4218, 4285, 4341, 4404, 4467, 4527, 4584, 4631, 4689, 4725]}, {"subject_area"=>"/Biology and life sciences/Molecular biology", "average_usage"=>[334, 613, 751, 863, 964, 1078, 1181, 1270, 1358, 1443, 1536, 1616, 1690, 1768, 1826, 1902, 1969, 2031, 2095, 2149, 2202, 2258, 2305, 2349, 2417, 2482, 2541, 2598, 2662, 2720, 2768, 2832, 2897, 2947, 2998, 3058, 3121, 3175, 3235, 3292, 3368, 3442, 3507, 3563, 3621, 3676, 3736, 3802, 3852, 3922, 3977, 4025, 4071, 4141, 4199, 4258, 4316, 4376, 4422, 4467, 4518]}, {"subject_area"=>"/Medicine and health sciences/Pulmonology", "average_usage"=>[355, 609, 758, 894, 995, 1098, 1194, 1269, 1352, 1425, 1519, 1632, 1716, 1798, 1929, 2031, 2117, 2185, 2252, 2315, 2387, 2455, 2532, 2616, 2736, 2798, 2845, 2901, 2957, 3022, 3111, 3159, 3241, 3294, 3341, 3373, 3408, 3445, 3506, 3555, 3619, 3686, 3745, 3826, 3894, 3961, 4015, 4055, 4120, 4165, 4218, 4265, 4315, 4380, 4435, 4500, 4539, 4582, 4635, 4703, 4740, 4753]}, {"subject_area"=>"/People and places", "average_usage"=>[340, 621, 760, 868, 956, 1048, 1152, 1251, 1334, 1414, 1499, 1591, 1663, 1753, 1823, 1893, 1955, 2026, 2094, 2178, 2255, 2327, 2390, 2467, 2535, 2613, 2680, 2731, 2780, 2851, 2909, 2986, 3049, 3107, 3177, 3238, 3292, 3360, 3437, 3503, 3564, 3622, 3710, 3812, 3860, 3932, 4002, 4075, 4112, 4156, 4247, 4296, 4386, 4455, 4550, 4613, 4662, 4694, 4869, 4937, 4982]}, {"subject_area"=>"/People and places/Geographical locations", "average_usage"=>[300, 591, 704, 817, 886, 1024, 1162, 1246, 1348, 1434, 1532, 1638, 1710, 1789, 1862, 1949, 2030, 2092, 2161, 2215, 2278, 2341, 2414, 2486, 2553, 2609, 2671, 2749, 2796, 2878, 2988, 3035, 3107, 3157, 3210, 3281, 3381, 3421, 3485, 3598, 3691, 3776, 3836, 3913, 3999, 4047, 4095, 4156, 4219, 4285, 4359, 4424, 4510, 4588, 4647, 4722, 4808, 4855, 4904, 4967, 5011, 5041]}]}

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