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We combined bioinformatics analysis and functional screening in three cell lines to provide evidence for regulatory activity. We then performed a genetic study to test the association of our selected candidates with disease susceptibility followed by the functional validation of the risk haplotype.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      We prioritized two\n                      <jats:italic>DPP9<\/jats:italic>\n                      variants within a haplotype that increases the risk of severe COVID-19. This haplotype exhibited increased regulatory activity and altered transcription factor binding, suggesting its role in influencing COVID-19 severity by modulating DPP9 expression in immune and lung cell types.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>The interest of our study lies in the functional characterization of regulatory variants responsible for increased levels of DPP9 and lung damage observed in patients with severe COVID-19. These findings advance our understanding of genetic risk factors for COVID-19 and highlight functional SNPs that may guide future therapeutic research.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s13073-026-01703-0","type":"journal-article","created":{"date-parts":[[2026,7,2]],"date-time":"2026-07-02T16:54:13Z","timestamp":1783011253000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Identification and functional characterization of regulatory variants in DPP9 associated with COVID-19 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The study is sponsored by Institut Pasteur (Pasteur ID-RCB Number: 2012-A00238-35), and was conducted as a single centre interventional study without an investigational product. The original protocol was registered under ClinicalTrials.gov (study# NCT01699893). The samples and data used in this study were formally established as the Milieu Interieur biocollection (NCT03905993), with approvals by the Comit\u00e9 de Protection des Personnes \u2013 Sud M\u00e9diterran\u00e9e and the Commission nationale de l'informatique et des libert\u00e9s (CNIL) on April 11, 2018, in accordance with the Declarations of Helsinki. All participants gave informed written consent.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"138"}}