{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T19:40:14Z","timestamp":1790278814288,"version":"4.1.0"},"reference-count":49,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,8,6]],"date-time":"2019-08-06T00:00:00Z","timestamp":1565049600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2019,8,6]],"date-time":"2019-08-06T00:00:00Z","timestamp":1565049600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Thalidomide is widely used for several diseases; however, it causes malformations in embryos exposed during pregnancy. The complete understanding of the mechanisms by which thalidomide affects the embryo development has not yet been obtained. The phenotypic similarity makes TE a phenocopy of syndromes caused by mutations in <jats:italic>ESCO2<\/jats:italic>, <jats:italic>SALL4<\/jats:italic> and <jats:italic>TBX5<\/jats:italic> genes. Recently, SALL4 and TBX5 were demonstrated to be thalidomide targets. To understand if these genes act in the TE development, we sequenced them in 27 individuals with TE; we verified how thalidomide affect them in human pluripotent stem cells (hPSCs) through a differential gene expression (DGE) analysis from GSE63935; and we evaluated how these genes are functionally related through an interaction network analysis. We identified 8 variants in <jats:italic>ESCO2<\/jats:italic>, 15 in <jats:italic>SALL4<\/jats:italic> and 15 in <jats:italic>TBX5<\/jats:italic>. We compared allelic frequencies with data from ExAC, 1000 Genomes and ABraOM databases; eight variants were significantly different (p\u2009&lt;\u20090.05). Eleven variants in <jats:italic>SALL4<\/jats:italic> and <jats:italic>TBX5<\/jats:italic> were previously associated with cardiac diseases or malformations; however, in TE sample there was no association. Variant effect prediction tools showed 97% of the variants with potential to influence in these genes regulation. DGE analysis showed a significant reduction of <jats:italic>ESCO2<\/jats:italic> in hPSCs after thalidomide exposure.<\/jats:p>","DOI":"10.1038\/s41598-019-47739-8","type":"journal-article","created":{"date-parts":[[2019,8,6]],"date-time":"2019-08-06T10:22:49Z","timestamp":1565086969000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["The role of ESCO2, SALL4 and TBX5 genes in the susceptibility to thalidomide teratogenesis"],"prefix":"10.1038","volume":"9","author":[{"given":"Julia do Amaral","family":"Gomes","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thayne Woycinck","family":"Kowalski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lucas Rosa","family":"Fraga","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabriel S.","family":"Macedo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Teresa Vieira","family":"Sanseverino","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lav\u00ednia","family":"Schuler-Faccini","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6339-4869","authenticated-orcid":false,"given":"Fernanda Sales Luiz","family":"Vianna","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,8,6]]},"reference":[{"key":"47739_CR1","doi-asserted-by":"publisher","first-page":"140","DOI":"10.1002\/bdrc.21096","volume":"105","author":"N Vargesson","year":"2015","unstructured":"Vargesson, N. Thalidomide-induced teratogenesis: history and mechanisms. Birth Defects Res C Embryo Today 105, 140\u2013156 (2015).","journal-title":"Birth Defects Res C Embryo Today"},{"key":"47739_CR2","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1084\/jem.173.3.699","volume":"173","author":"EP Sampaio","year":"1991","unstructured":"Sampaio, E. P., Sarno, E. N., Galilly, R., Cohn, Z. A. & Kaplan, G. Thalidomide selectively inhibits tumor necrosis factor alpha production by stimulated human monocytes. J Exp Med 173, 699\u2013703 (1991).","journal-title":"J Exp Med"},{"key":"47739_CR3","doi-asserted-by":"publisher","first-page":"4082","DOI":"10.1073\/pnas.91.9.4082","volume":"91","author":"RJ D\u2019Amato","year":"1994","unstructured":"D\u2019Amato, R. J., Loughnan, M. S., Flynn, E. & Folkman, J. Thalidomide is an inhibitor of angiogenesis. Proc Natl Acad Sci USA 91, 4082\u20134085 (1994).","journal-title":"Proc Natl Acad Sci USA"},{"key":"47739_CR4","doi-asserted-by":"publisher","first-page":"3401","DOI":"10.1590\/S1413-81232013001100029","volume":"18","author":"FJ Paumgartten","year":"2013","unstructured":"Paumgartten, F. J. & de Souza, N. R. Clinical use and control of the dispensing of thalidomide in Brasilia-Federal District, Brazil, from 2001 to 2012. Cien Saude Cole 18, 3401\u20133048 (2013).","journal-title":"Cien Saude Cole"},{"key":"47739_CR5","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/j.reprotox.2015.03.007","volume":"53","author":"F Sales Luiz Vianna","year":"2015","unstructured":"Sales Luiz Vianna, F. et al. Pharmacoepidemiology and thalidomide embryopathy surveillance in Brazil. Reprod Toxicol 53, 63\u201367 (2015).","journal-title":"Reprod Toxicol"},{"key":"47739_CR6","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.ejmg.2016.09.015","volume":"60","author":"VF Sales Luiz","year":"2017","unstructured":"Sales Luiz, V. F., Kowalski, T. W., Fraga, L. R., Sanseverino, M. T. & Schuler-Faccini, L. The impact of thalidomide use in birth defects in Brazil. Eur J Med Genet 60, 12\u201315 (2017).","journal-title":"Eur J Med Genet"},{"key":"47739_CR7","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1097\/MCD.0b013e32835ffc58","volume":"22","author":"FS Vianna","year":"2013","unstructured":"Vianna, F. S. et al. Recognition of the phenotype of thalidomide embryopathy in countries endemic for leprosy: new cases and review of the main dysmorphological findings. Clin Dysmorphol 22, 59\u201363 (2013).","journal-title":"Clin Dysmorphol"},{"key":"47739_CR8","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1016\/S0095-5108(18)30810-8","volume":"13","author":"CG Newman","year":"1986","unstructured":"Newman, C. G. The thalidomide syndrome: risks of exposure and spectrum of malformations. Clin Perinatol 13, 555\u2013573 (1986).","journal-title":"Clin Perinatol"},{"key":"47739_CR9","doi-asserted-by":"publisher","first-page":"21735","DOI":"10.1371\/journal.pone.0021735","volume":"6","author":"FS Vianna","year":"2011","unstructured":"Vianna, F. S. et al. Epidemiological surveillance of birth defects compatible with thalidomide embryopathy in Brazil. PLoS One 6, 21735 (2011).","journal-title":"PLoS One"},{"key":"47739_CR10","doi-asserted-by":"publisher","first-page":"186","DOI":"10.1016\/j.reprotox.2012.05.004","volume":"34","author":"M Cassina","year":"2012","unstructured":"Cassina, M., Salviati, L., Di Gianantonio, E. & Clementi, M. Genetic susceptibility to teratogens: state of the art. Reprod Toxicol 34, 186\u2013191 (2012).","journal-title":"Reprod Toxicol"},{"key":"47739_CR11","doi-asserted-by":"publisher","DOI":"10.1038\/srep23404","volume":"6","author":"TW Kowalski","year":"2016","unstructured":"Kowalski, T. W. et al. New Findings in eNOS gene and Thalidomide Embryopathy Suggest pre-transcriptional effect variants as susceptibility factors. Sci Rep 6, 23404 (2016).","journal-title":"Sci Rep"},{"key":"47739_CR12","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/j.reprotox.2016.10.003","volume":"66","author":"FS Vianna","year":"2016","unstructured":"Vianna, F. S. et al. Genomic and in silico analyses of CRBN gene and thalidomide embryopathy in humans. Reprod Toxicol 66, 99\u2013106 (2016).","journal-title":"Reprod Toxicol"},{"key":"47739_CR13","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1136\/jmg.10.1.34","volume":"10","author":"W Lenz","year":"1973","unstructured":"Lenz, W. Phenocopies. J Med Genet 10, 34\u201349 (1973).","journal-title":"J Med Genet"},{"key":"47739_CR14","doi-asserted-by":"publisher","first-page":"1117","DOI":"10.1086\/498695","volume":"77","author":"B Sch\u00fcle","year":"2005","unstructured":"Sch\u00fcle, B., Oviedo, A., Johnston, K., Pai, S. & Francke, U. Inactivating mutations in ESCO2 cause SC phocomelia and Roberts syndrome: no phenotype-genotype correlation. Am J Hum Genet 77, 1117\u20131128 (2005).","journal-title":"Am J Hum Genet"},{"key":"47739_CR15","doi-asserted-by":"publisher","first-page":"468","DOI":"10.1038\/ng1548","volume":"37","author":"H Vega","year":"2005","unstructured":"Vega, H. et al. Roberts syndrome is caused by mutations in ESCO2, a human homolog of yeast ECO1 that is essential for the establishment of sister chromatid cohesion. Nat Genet 37, 468\u2013470 (2005).","journal-title":"Nat Genet"},{"key":"47739_CR16","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1038\/ng0197-21","volume":"15","author":"QY Li","year":"1997","unstructured":"Li, Q. Y. et al. Holt-Oram syndrome is caused by mutations in TBX5, a member of the Brachyury (T) gene family. Nat Genet 15, 21\u201329 (1997).","journal-title":"Nat Genet"},{"key":"47739_CR17","doi-asserted-by":"publisher","first-page":"1195","DOI":"10.1086\/343821","volume":"71","author":"R Al-Baradie","year":"2002","unstructured":"Al-Baradie, R. et al. Duane radial ray syndrome (Okihiro syndrome) maps to 20q13 and results from mutations in SALL4, a new member of the SAL family. Am J Hum Genet 71, 1195\u20131199 (2002).","journal-title":"Am J Hum Genet"},{"key":"47739_CR18","doi-asserted-by":"publisher","first-page":"1165","DOI":"10.1242\/dev.02259","volume":"133","author":"MP Harvey","year":"2006","unstructured":"Harvey, M. P. & Logan, O. Sall4 acts downstream of tbx5 and is required for pectoral fin outgrowth. Development 133, 1165\u20131173 (2006).","journal-title":"Development"},{"key":"47739_CR19","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1038\/ng1707","volume":"38","author":"K Koshiba-Takeuchi","year":"2006","unstructured":"Koshiba-Takeuchi, K. et al. Cooperative and antagonistic interactions between Sall4 and Tbx5 pattern the mouse limb and heart. Nat Genet 38, 175\u2013183 (2006).","journal-title":"Nat Genet"},{"key":"47739_CR20","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.ejmg.2016.09.011","volume":"60","author":"M Cassina","year":"2017","unstructured":"Cassina, M., Cagnoli, G. A., Zuccarello, D., Di Gianantonio, E. & Clementi, M. Human teratogens and genetic phenocopies: Understanding pathogenesis through human genes mutation. Eur J Med Genet 60, 22\u201331 (2017).","journal-title":"Eur J Med Genet"},{"key":"47739_CR21","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.4161\/cc.7.9.5793","volume":"7","author":"J Knobloch","year":"2008","unstructured":"Knobloch, J. & R\u00fcther, U. Shedding light on an old mystery: thalidomide suppresses survival pathways to induce limb defects. Cell Cycle 7, 1121\u20131127 (2008).","journal-title":"Cell Cycle"},{"key":"47739_CR22","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-01641-3","volume":"7","author":"A Khalil","year":"2017","unstructured":"Khalil, A. et al. A HAND to TBX5 Explains the Link Between Thalidomide and Cardiac Diseases. Sci Rep 7, 1416 (2017).","journal-title":"Sci Rep"},{"key":"47739_CR23","doi-asserted-by":"publisher","first-page":"e38430","DOI":"10.7554\/eLife.38430","volume":"7","author":"KA Donovan","year":"2018","unstructured":"Donovan, K. A. et al. Thalidomide promotes degradation of SALL4, a transcription factor implicated in Duane Radial Ray Syndrome. Elife 7, e38430 (2018).","journal-title":"Elife"},{"key":"47739_CR24","doi-asserted-by":"publisher","first-page":"981","DOI":"10.1038\/s41589-018-0129-x","volume":"14","author":"ME Matyskiela","year":"2018","unstructured":"Matyskiela, M. E. et al. SALL4 mediates teratogenicity as a thalidomide-dependent cereblon substrate. Nat Chem Biol 14, 981\u2013987 (2018).","journal-title":"Nat Chem Biol"},{"key":"47739_CR25","doi-asserted-by":"publisher","first-page":"980","DOI":"10.1093\/nar\/gkt1113","volume":"42","author":"MJ Landrum","year":"2014","unstructured":"Landrum, M. J. et al. ClinVar: public archive of relationships among sequence variation and human phenotype. Nucleic Acids Res 42, 980\u2013D985 (2014).","journal-title":"Nucleic Acids Res"},{"key":"47739_CR26","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1038\/gim.2015.30","volume":"17","author":"S Richards","year":"2015","unstructured":"Richards, S. et al. ACMG Laboratory Quality Assurance Committee. Standards and guidelines for the interpretation of sequence variants: a joint consensus recommendation of the American College of Medical Genetics and Genomics and the Association for Molecular Pathology. Genet Med 17, 405\u2013424 (2015).","journal-title":"Genet Med"},{"key":"47739_CR27","doi-asserted-by":"publisher","first-page":"12516","DOI":"10.1073\/pnas.1516645112","volume":"112","author":"MP Schwartz","year":"2015","unstructured":"Schwartz, M. P. et al. Human pluripotent stem cell-derived neural constructs for predicting neural toxicity. Proc Natl Acad Sci USA 112, 12516\u201312521 (2015).","journal-title":"Proc Natl Acad Sci USA"},{"key":"47739_CR28","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1007\/s00109-004-0603-7","volume":"83","author":"JC Knight","year":"2005","unstructured":"Knight, J. C. Regulatory polymorphisms underlying complex disease traits. J Mol Med (Berl) 83, 97\u2013109 (2005).","journal-title":"J Mol Med (Berl)"},{"key":"47739_CR29","doi-asserted-by":"publisher","first-page":"592","DOI":"10.1016\/j.ymgme.2012.09.020","volume":"107","author":"PJ Lupo","year":"2012","unstructured":"Lupo, P. J. et al. A GCH1 haplotype and risk of neural tube defects in the National Birth Defects Prevention Study. Mol Genet Metab 107, 592\u2013595 (2012).","journal-title":"Mol Genet Metab"},{"key":"47739_CR30","doi-asserted-by":"publisher","first-page":"610","DOI":"10.1101\/gr.7179508","volume":"18","author":"RD Morin","year":"2008","unstructured":"Morin, R. D. et al. Application of massively parallel sequencing to microRNA profiling and discovery in human embryonic stem cells. Genome Res 18, 610\u2013621 (2008).","journal-title":"Genome Res"},{"key":"47739_CR31","doi-asserted-by":"publisher","first-page":"1345","DOI":"10.1126\/science.1177319","volume":"327","author":"T Ito","year":"2010","unstructured":"Ito, T. et al. Identification of a primary target of thalidomide teratogenicity. Science 327, 1345\u20131350 (2010).","journal-title":"Science"},{"key":"47739_CR32","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1002\/(SICI)1096-9926(199911)60:5<306::AID-TERA11>3.0.CO;2-Y","volume":"60","author":"MT Miller","year":"1999","unstructured":"Miller, M. T. & Str\u00f6mland, K. Teratogen update: Thalidomide: A review, with a focus on ocular findings and new potential uses. Teratology 60, 306\u2013321 (1999).","journal-title":"Teratology"},{"key":"47739_CR33","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/bs.ctdb.2016.08.008","volume":"122","author":"JD Steimle","year":"2017","unstructured":"Steimle, J. D. & Moskowitz, I. P. TBX5: A Key Regulator of Heart Development. Curr Top Dev Biol 122, 195\u2013221 (2017).","journal-title":"Curr Top Dev Biol"},{"key":"47739_CR34","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1038\/ng.511","volume":"42","author":"H Holm","year":"2010","unstructured":"Holm, H. et al. Several common variants modulate heart rate, PR interval and QRS duration. Nat Genet 42, 117\u2013122 (2010).","journal-title":"Nat Genet"},{"key":"47739_CR35","doi-asserted-by":"crossref","first-page":"e001978","DOI":"10.1161\/CIRCGEN.117.001978","volume":"11","author":"JO Szot","year":"2010","unstructured":"Szot, J. O. et al. A Screening Approach to Identify Clinically Actionable Variants Causing Congenital Heart Disease in Exome. Data. Circ Genom Precis Med 11, e001978 (2010).","journal-title":"Data. Circ Genom Precis Med"},{"key":"47739_CR36","doi-asserted-by":"publisher","first-page":"e57216","DOI":"10.1371\/journal.pone.0057216","volume":"8","author":"R Pazoki","year":"2013","unstructured":"Pazoki, R. et al. SNPs Identified as Modulators of ECG Traits in the General Population Do Not Markedly Affect ECG Traits during Acute Myocardial Infarction nor Ventricular Fibrillation Risk in This Condition. PLoS One 8, e57216 (2013).","journal-title":"PLoS One"},{"key":"47739_CR37","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1186\/1755-8794-7-56","volume":"7","author":"EM Bonachea","year":"2014","unstructured":"Bonachea, E. M. et al. Use of a targeted, combinatorial next-generation sequencing approach for the study of bicuspid aortic valve. BMC Medical Genomic 7, 56 (2014).","journal-title":"BMC Medical Genomic"},{"key":"47739_CR38","doi-asserted-by":"publisher","first-page":"17026","DOI":"10.1038\/celldisc.2017.26","volume":"3","author":"F Wang","year":"2017","unstructured":"Wang, F. et al. A TBX5 3\u2032UTR variant increases the risk of congenital heart disease in the Han Chinese population. Cell Discov 3, 17026 (2017).","journal-title":"Cell Discov"},{"key":"47739_CR39","doi-asserted-by":"publisher","first-page":"5823","DOI":"10.3892\/mmr.2017.7334","volume":"16","author":"Y Yamada","year":"2017","unstructured":"Yamada, Y. et al. Identification of TNFSF13, SPATC1L, SLC22A25 and SALL4 as novel susceptibility loci for atrial fibrillation by an exome-wide association study. Mol Med Rep 16, 5823\u20135832 (2017).","journal-title":"Mol Med Rep"},{"key":"47739_CR40","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1002\/1097-0177(200009)219:1<90::AID-DVDY1033>3.0.CO;2-L","volume":"219","author":"CJ Hatcher","year":"2000","unstructured":"Hatcher, C. J., Goldstein, M. M., Mah, C. S., Delia, C. S. & Basson, C. T. Identification and localization of TBX5 transcription factor during human cardiac morphogenesis. Dev Dyn 219, 90\u20135 (2000).","journal-title":"Dev Dyn"},{"key":"47739_CR41","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.gene.2016.02.019","volume":"584","author":"H Tatetsu","year":"2016","unstructured":"Tatetsu, H. et al. SALL4, the missing link between stem cells, development and cancer. Gene 584, 111\u20139 (2016).","journal-title":"Gene"},{"key":"47739_CR42","doi-asserted-by":"publisher","first-page":"258","DOI":"10.1093\/nar\/gkg034","volume":"31","author":"C von Mering","year":"2003","unstructured":"von Mering, C. et al. STRING: a database of predicted functional association between proteins. Nucleic Acids Res 31, 258\u2013261 (2003).","journal-title":"Nucleic Acids Res"},{"key":"47739_CR43","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1089\/152308604771978291","volume":"6","author":"JM Hansen","year":"2004","unstructured":"Hansen, J. M. & Harris, C. A novel hypothesis for thalidomide-induced limb teratogenesis: redox misregulation of the NF-kappaB pathway. Antioxid Redox Signal 6, 1\u201314 (2004).","journal-title":"Antioxid Redox Signal"},{"key":"47739_CR44","doi-asserted-by":"publisher","first-page":"794","DOI":"10.1002\/bdra.23376","volume":"103","author":"TW Kowalski","year":"2015","unstructured":"Kowalski, T. W., Sanseverino, M. T., Schuler-Faccini, L. & Vianna, F. S. Thalidomide embryopathy: Follow-up of cases born between 1959 and 2010. Birth Defects Res A Clin Mol Teratol 103, 794\u2013803 (2015).","journal-title":"Birth Defects Res A Clin Mol Teratol"},{"key":"47739_CR45","unstructured":"Gordillo, M., Vega, H. & Jabs, E. W. Roberts Syndrome in GeneReviews (R). Seattle, WA: in press (2013)."},{"key":"47739_CR46","unstructured":"Kohlhase, J. SALL4-Related Disorders in GeneReviews (R). Seattle, WA: in press (2015)."},{"key":"47739_CR47","unstructured":"McDermott, D. A., Fong, J. C. & Basson, C. T. Holt-Oram Syndrome in GeneReviews (R). Seattle, WA: in press (2015)."},{"key":"47739_CR48","doi-asserted-by":"publisher","first-page":"5075","DOI":"10.1073\/pnas.1421949112","volume":"112","author":"R Akiyama","year":"2015","unstructured":"Akiyama, R. et al. Sall4-Gli3 system in early limb progenitors is essential for the development of limb skeletal elements. Proc Natl Acad Sci USA 112, 5075\u20135080 (2015).","journal-title":"Proc Natl Acad Sci USA"},{"key":"47739_CR49","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/bs.ctdb.2016.08.009","volume":"122","author":"CJ Sheeba","year":"2017","unstructured":"Sheeba, C. J. & Logan, M. P. The Roles of T-Box Genes in Vertebrate Limb Development. Curr Top Dev Biol 122, 355\u2013381 (2017).","journal-title":"Curr Top Dev Biol"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-47739-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-47739-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-47739-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,17]],"date-time":"2022-12-17T12:21:38Z","timestamp":1671279698000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-47739-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,8,6]]},"references-count":49,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,12]]}},"alternative-id":["47739"],"URL":"https:\/\/doi.org\/10.1038\/s41598-019-47739-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,8,6]]},"assertion":[{"value":"11 January 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 June 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 August 2019","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"11413"}}