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<jats:title>INTRODUCTION<\/jats:title>\n                    <jats:p>\n                      An extra copy of the amyloid precursor protein (\n                      <jats:italic>APP<\/jats:italic>\n                      ) gene causes autosomal dominant Alzheimer's disease (AD) and AD in Down syndrome (DS), but the factors underlying variability in age at onset (AAO) remain unclear. We investigated whether sporadic AD risk variants modify AAO.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>METHODS<\/jats:title>\n                    <jats:p>\n                      We analyzed clinical and genetic data from 100\n                      <jats:italic>APP<\/jats:italic>\n                      duplication (\n                      <jats:italic>APP<\/jats:italic>\n                      dup) carriers and 957 individuals with DS. Cox models assessed associations of apolipoprotein E (\n                      <jats:italic>APOE<\/jats:italic>\n                      ) \u03b52 and \u03b54 and the AD genetic risk score (AD\u2010GRS; excluding\n                      <jats:italic>APOE<\/jats:italic>\n                      and chromosome 21 variants) with AAO.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>RESULTS<\/jats:title>\n                    <jats:p>\n                      Mean AAO was earlier in\n                      <jats:italic>APP<\/jats:italic>\n                      dup than DS (51\u00a0\u00b1\u00a07\u00a0vs. 53\u00a0\u00b1\u00a06 years;\n                      <jats:italic>P\u00a0=<\/jats:italic>\n                      \u00a00.0005).\n                      <jats:italic>APOE<\/jats:italic>\n                      \u03b52 delayed onset (hazard ratio [HR]\u00a0=\u00a00.47,\n                      <jats:italic>P\u00a0&lt;<\/jats:italic>\n                      \u00a00.0001), whereas\n                      <jats:italic>APOE<\/jats:italic>\n                      \u03b54 (HR\u00a0=\u00a01.5,\n                      <jats:italic>P\u00a0=<\/jats:italic>\n                      \u00a00.0003) and higher AD\u2010GRS (HR\u00a0=\u00a01.3 per standard deviation,\n                      <jats:italic>P\u00a0&lt;<\/jats:italic>\n                      \u00a00.0001) accelerated onset. Predicted median AAO differed by 10 years between lowest and highest genetic risk.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>DISCUSSION<\/jats:title>\n                    <jats:p>\n                      Sporadic AD genetic risk factors are important modifiers of AAO in\n                      <jats:italic>APP<\/jats:italic>\n                      dup and DS, explaining part of the marked variability in onset.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1002\/alz.71738","type":"journal-article","created":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T09:47:46Z","timestamp":1786182466000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["<i>APOE<\/i>\n                    and genetic risk variants influence Alzheimer's disease onset in carriers of an extra copy of\n                    <i>APP<\/i>\n                    , with and without Down syndrome"],"prefix":"10.1002","volume":"22","author":[{"given":"Joan","family":"Groeneveld","sequence":"first","affiliation":[{"name":"Genomics of Neurodegenerative Diseases and Aging, Human Genetics, Amsterdam UMC location VUmc Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Danna","family":"Perlaza","sequence":"additional","affiliation":[{"name":"Sant Pau Memory Unit, IR SANT PAU Hospital de la Santa Creu i Sant Pau  Barcelona Spain"},{"name":"Network Center for Biomedical Research in Neurodegenerative Diseases (CIBERNED)  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cl\u00e0udia","family":"Oliv\u00e9","sequence":"additional","affiliation":[{"name":"Ace Alzheimer Center Barcelona Universitat Internacional de Catalunya (UIC)  Barcelona Spain"},{"name":"Doctorate in Biotechnology, Facultat de Farm\u00e0cia i Ci\u00e8ncies de l'Alimentaci\u00f3 Universitat de Barcelona  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lou","family":"Grangeon","sequence":"additional","affiliation":[{"name":"Inserm U1245 and, Department of Neurology and CNRMAJ Univ Rouen Normandie, Normandie Univ, CHU Rouen  Rouen France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Niccolo","family":"Tesi","sequence":"additional","affiliation":[{"name":"Genomics of Neurodegenerative Diseases and Aging, Human Genetics, Amsterdam UMC location VUmc Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Delft Bioinformatics Lab Delft University of Technology  Delft the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aude","family":"Nicolas","sequence":"additional","affiliation":[{"name":"Inserm, U1167\u2010RID\u2010AGE Facteurs de risque et d\u00e9terminants mol\u00e9culaires des maladies li\u00e9es au vieillissement Univ. Lille, CHU Lille, Institut Pasteur de Lille  Lille France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenyang","family":"Jiang","sequence":"additional","affiliation":[{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Itziar","family":"de Rojas","sequence":"additional","affiliation":[{"name":"Network Center for Biomedical Research in Neurodegenerative Diseases (CIBERNED)  Madrid Spain"},{"name":"Ace Alzheimer Center Barcelona Universitat Internacional de Catalunya (UIC)  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David","family":"Wallon","sequence":"additional","affiliation":[{"name":"Inserm U1245 and, Department of Neurology and CNRMAJ Univ Rouen Normandie, Normandie Univ, CHU Rouen  Rouen France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"St\u00e9phane","family":"Rousseau","sequence":"additional","affiliation":[{"name":"Univ Rouen Normandie, Normandie Univ, CHU Rouen Inserm U1245 and Department of Genetics and CNRMAJ  Rouen France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marta","family":"Rovira","sequence":"additional","affiliation":[{"name":"Ace Alzheimer Center Barcelona Universitat Internacional de Catalunya (UIC)  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Diego Real","family":"de As\u00faa","sequence":"additional","affiliation":[{"name":"Adult Down Syndrome Unit, Department of Internal Medicine Hospital Universitario de La Princesa Instituto de Investigaci\u00f3n Sanitaria Hospital Universitario de La Princesa  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fernando","family":"Moldenhauer","sequence":"additional","affiliation":[{"name":"Adult Down Syndrome Unit, Department of Internal Medicine Hospital Universitario de La Princesa Instituto de Investigaci\u00f3n Sanitaria Hospital Universitario de La Princesa  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruihao","family":"Mu","sequence":"additional","affiliation":[{"name":"Instituto de Investigaci\u00f3n Sanitaria Hospital Universitario de La Princesa  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K\u00e9vin","family":"Cassinari","sequence":"additional","affiliation":[{"name":"Inserm U1245 and, Department of Neurology and CNRMAJ Univ Rouen Normandie, Normandie Univ, CHU Rouen  Rouen France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aline","family":"Zarea","sequence":"additional","affiliation":[{"name":"Inserm U1245 and, Department of Neurology and CNRMAJ Univ Rouen Normandie, Normandie Univ, CHU Rouen  Rouen France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joaquim Aumatell","family":"Escabias","sequence":"additional","affiliation":[{"name":"Sant Pau Memory Unit, IR SANT PAU Hospital de la Santa Creu i Sant Pau  Barcelona Spain"},{"name":"Network Center for Biomedical Research in Neurodegenerative Diseases (CIBERNED)  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jean\u2010Charles","family":"Lambert","sequence":"additional","affiliation":[{"name":"Inserm, U1167\u2010RID\u2010AGE Facteurs de risque et d\u00e9terminants mol\u00e9culaires des maladies li\u00e9es au vieillissement Univ. Lille, CHU Lille, Institut Pasteur de Lille  Lille France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yolande A. L.","family":"Pijnenburg","sequence":"additional","affiliation":[{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marc","family":"Hulsman","sequence":"additional","affiliation":[{"name":"Genomics of Neurodegenerative Diseases and Aging, Human Genetics, Amsterdam UMC location VUmc Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Everard G. B.","family":"Vijverberg","sequence":"additional","affiliation":[{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johannes","family":"Levin","sequence":"additional","affiliation":[{"name":"Department of Neurology Ludwig\u2010Maximilians\u2010Universit\u00e4t M\u00fcnchen  Munich Germany"},{"name":"Site Munich German Center for Neurodegenerative Diseases  Munich Germany"},{"name":"Munich Cluster for Systems Neurology (SyNergy)  Munich Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mathias","family":"Jucker","sequence":"additional","affiliation":[{"name":"German Center for Neurodegenerative Diseases (DZNE)  T\u00fcbingen Germany"},{"name":"Hertie Institute for Clinical Brain Research University of T\u00fcbingen  T\u00fcbingen Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eric","family":"McDade","sequence":"additional","affiliation":[{"name":"Department of Neurology Washington University School of Medicine  St. Louis Missouri USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan","family":"Fortea","sequence":"additional","affiliation":[{"name":"Sant Pau Memory Unit, IR SANT PAU Hospital de la Santa Creu i Sant Pau  Barcelona Spain"},{"name":"Network Center for Biomedical Research in Neurodegenerative Diseases (CIBERNED)  Madrid Spain"},{"name":"Barcelona Down Medical Center Fundaci\u00f3 Catalana S\u00edndrome de Down  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Henne","family":"Holstege","sequence":"additional","affiliation":[{"name":"Genomics of Neurodegenerative Diseases and Aging, Human Genetics, Amsterdam UMC location VUmc Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"},{"name":"VIB Center for Brain and Disease Research  Leuven Belgium"},{"name":"Department of Neurosciences Leuven Brain Institute KU Leuven  Leuven Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Flora H.","family":"Duits","sequence":"additional","affiliation":[{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"},{"name":"Neurochemistry Laboratory Department of Laboratory Medicine, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lisa","family":"Vermunt","sequence":"additional","affiliation":[{"name":"Alzheimer Center Amsterdam, Neurology, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"},{"name":"Neurodegeneration Amsterdam Neuroscience  Amsterdam the Netherlands"},{"name":"Neurochemistry Laboratory Department of Laboratory Medicine, Amsterdam UMC Vrije Universiteit Amsterdam  Amsterdam the Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maulikkumar","family":"Patel","sequence":"additional","affiliation":[{"name":"Department of Psychiatry Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Neurogenomics and Informatics Center Washington University School of Medicine  St. Louis Missouri USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthew","family":"Johnson","sequence":"additional","affiliation":[{"name":"Department of Psychiatry Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Neurogenomics and Informatics Center Washington University School of Medicine  St. Louis Missouri USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alan E.","family":"Renton","sequence":"additional","affiliation":[{"name":"Ronald M. 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Loeb Center for Alzheimer's Disease Icahn School of Medicine at Mount Sinai  New York New York USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carlos","family":"Cruchaga","sequence":"additional","affiliation":[{"name":"Department of Neurology Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Department of Psychiatry Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Neurogenomics and Informatics Center Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Hope Center for Neurological Disorders Washington University School of Medicine  St. Louis Missouri USA"},{"name":"Knight Alzheimer Disease Research Center Washington University School of Medicine  St. Louis Missouri USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cyril","family":"Pottier","sequence":"additional","affiliation":[{"name":"Department of Neurology Washington University School of Medicine  St. Louis Missouri 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