{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T05:19:07Z","timestamp":1787894347948,"version":"build-2784847793"},"reference-count":32,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T00:00:00Z","timestamp":1786147200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T00:00:00Z","timestamp":1786147200000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["AG066206"],"award-info":[{"award-number":["AG066206"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["GM127063"],"award-info":[{"award-number":["GM127063"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["AG066515"],"award-info":[{"award-number":["AG066515"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Genetic Epidemiology"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Genome\u2010wide association studies (GWASs) have been extensively adopted to depict the underlying genetic architecture of complex traits. Recent studies show that knockoff\u2010based methods can identify variants with unique, potentially causal effects on phenotypes. However, their statistical validity and effectiveness in studies with related individuals, such as the UK Biobank, remain unexplored. In this paper, we extensively evaluate a simple and effective analytical strategy that integrates GhostKnockoffs and state\u2010of\u2010the\u2010art marginal association tests. We show that this approach is robust to arbitrary relatedness structure as long as the input\n                    <jats:italic>Z<\/jats:italic>\n                    \u2010scores are derived from valid generalized linear mixed models. This robustness also extends GhostKnockoffs to other GWASs settings, including meta\u2010analysis of studies with sample overlap when the input score test\n                    <jats:italic>Z<\/jats:italic>\n                    \u2010scores are properly calibrated, and association test statistics beyond score tests in independent sample settings. We demonstrate the method's validity and practical utility using simulation studies and a meta\u2010analysis of nine European ancestral genome\u2010wide association studies and whole exome\/genome sequencing studies for the Alzheimer's disease.\n                  <\/jats:p>","DOI":"10.1002\/gepi.70051","type":"journal-article","created":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T11:35:13Z","timestamp":1786188913000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Robust Inference With Ghostknockoffs in Genome\u2010Wide Association Studies With Sample Relatedness"],"prefix":"10.1002","volume":"50","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5263-2419","authenticated-orcid":false,"given":"Xinran","family":"Qi","sequence":"first","affiliation":[{"name":"Department of Hematology and Hematopoietic Cell Transplantation City of Hope Comprehensive Cancer Center Duarte California USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7748-9033","authenticated-orcid":false,"given":"Michael E.","family":"Belloy","sequence":"additional","affiliation":[{"name":"Department of Neurology Washington University in Saint Louis Saint Louis Missouri USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8773-9558","authenticated-orcid":false,"given":"Jiaqi","family":"Gu","sequence":"additional","affiliation":[{"name":"Department of Mathematics and Statistics University of South Florida Tampa Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoxia","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Biomedical Informatics, Jacobs School of Medicine and Biomedical Sciences University at Buffalo Buffalo New York 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USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,8,8]]},"reference":[{"key":"e_1_2_11_2_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2007743117"},{"key":"e_1_2_11_3_1","doi-asserted-by":"publisher","DOI":"10.1212\/NXG.0000000000200012"},{"key":"e_1_2_11_4_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btv546"},{"key":"e_1_2_11_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380-018-0112-7"},{"key":"e_1_2_11_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2017.05.038"},{"key":"e_1_2_11_7_1","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.116.193987"},{"key":"e_1_2_11_8_1","doi-asserted-by":"publisher","DOI":"10.1111\/rssb.12265"},{"key":"e_1_2_11_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ajhg.2015.11.022"},{"key":"e_1_2_11_10_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-022-01087-y"},{"key":"e_1_2_11_11_1","doi-asserted-by":"publisher","DOI":"10.1186\/s13195-021-00808-5"},{"key":"e_1_2_11_12_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-34932-z"},{"key":"e_1_2_11_13_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-22889-4"},{"key":"e_1_2_11_14_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.4587"},{"key":"e_1_2_11_15_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-018-0311-9"},{"key":"e_1_2_11_16_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-019-0530-8"},{"key":"e_1_2_11_17_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2308-7"},{"key":"e_1_2_11_18_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-019-0358-2"},{"key":"e_1_2_11_19_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-020-0676-4"},{"key":"e_1_2_11_20_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-022-01225-6"},{"key":"e_1_2_11_21_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-022-01640-x"},{"key":"e_1_2_11_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ajhg.2009.11.001"},{"key":"e_1_2_11_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ajhg.2019.01.002"},{"key":"e_1_2_11_24_1","doi-asserted-by":"publisher","DOI":"10.1002\/zoo.1430050209"},{"key":"e_1_2_11_25_1","doi-asserted-by":"publisher","DOI":"10.1101\/gr.3709305"},{"key":"e_1_2_11_26_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-020-00776-w"},{"key":"e_1_2_11_27_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2105841118"},{"key":"e_1_2_11_28_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-14791-2"},{"key":"e_1_2_11_29_1","doi-asserted-by":"publisher","DOI":"10.1093\/biomet\/asy033"},{"key":"e_1_2_11_30_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkac1010"},{"key":"e_1_2_11_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ajhg.2011.05.029"},{"key":"e_1_2_11_32_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-021-04369-3"},{"key":"e_1_2_11_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ajhg.2022.08.013"}],"container-title":["Genetic 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