Meta-Analysis

. 2025 Jul 17;26(1):210.

doi: 10.1186/s13059-025-03564-z.

Multi-ancestry genome-wide meta-analysis of 56,241 individuals identifies known and novel cross-population and ancestry-specific associations as novel risk loci for Alzheimer's disease

Farid Rajabli^#^{1

2}, Penelope Benchek^#³, Giuseppe Tosto^#^{4

5}, Nicholas Kushch², Jin Sha⁶, Katrina Bazemore⁶, Congcong Zhu⁷, Wan-Ping Lee⁸, Jacob Haut⁶, Kara L Hamilton-Nelson², Nicholas R Wheeler^{3

9}, Yi Zhao⁸, John J Farrell⁷, Michelle A Grunin³, Yuk Yee Leung⁸, Pavel P Kuksa⁸, Donghe Li⁷, Eder Lucio da Fonseca², Jesse B Mez¹⁰, Ellen L Palmer³, Jagan Pillai¹¹, Richard M Sherva⁷, Yeunjoo E Song^{3

9}, Xiaoling Zhang^{7

12}, Takeshi Ikeuchi¹³, Taha Iqbal⁶, Omkar Pathak⁸, Otto Valladares⁸, Dolly Reyes-Dumeyer^{4

5}, Amanda B Kuzma⁸, Erin Abner¹⁴, Larry D Adams¹, Perrie M Adams¹⁵, Alyssa Aguirre¹⁶, Marilyn S Albert¹⁷, Roger L Albin^{18

19

20}, Mariet Allen²¹, Lisa Alvarez²², Liana G Apostolova^{23

24}, Steven E Arnold²⁵, Sanjay Asthana^{26

27

28}, Craig S Atwood^{26

27

28}, Sanford Auerbach¹⁰, Gayle Ayres¹⁶, Clinton T Baldwin⁷, Robert C Barber²², Lisa L Barnes^{29

30

31}, Sandra Barral^{4

5

32}, Thomas G Beach³³, James T Becker³⁴, Gary W Beecham², Duane Beekly³⁵, Bruno A Benitez³⁶, David Bennett^{29

31}, John Bertelson³⁷, Thomas D Bird^{38

39}, Deborah Blacker^{40

41}, Bradley F Boeve⁴², James D Bowen⁴³, Adam Boxer⁴⁴, James Brewer⁴⁵, James R Burke⁴⁶, Jeffrey M Burns⁴⁷, Joseph D Buxbaum^{48

49

50}, Nigel J Cairns⁵¹, Laura B Cantwell⁸, Chuanhai Cao⁵², Christopher S Carlson⁵³, Cynthia M Carlsson^{27

28}, Regina M Carney⁵⁴, Minerva M Carrasquillo²¹, Scott Chasse⁵⁵, Marie-Francoise Chesselet⁵⁶, Nathaniel A Chin^{26

27}, Helena C Chui⁵⁷, Jaeyoon Chung⁷, Suzanne Craft⁵⁸, Paul K Crane⁵⁹, David H Cribbs⁶⁰, Elizabeth A Crocco⁶¹, Carlos Cruchaga^{62

63}, Michael L Cuccaro^{1

2}, Munro Cullum¹⁵, Eveleen Darby⁶⁴, Barbara Davis⁶⁵, Philip L De Jager⁶⁶, Charles DeCarli⁶⁷, John DeToledo⁶⁸, Malcolm Dick⁶⁹, Dennis W Dickson²¹, Beth A Dombroski⁸, Rachelle S Doody⁶⁴, Ranjan Duara⁷⁰, NIlüfer Ertekin-Taner^{21

71}, Denis A Evans⁷², Kelley M Faber⁷³, Thomas J Fairchild⁷⁴, Kenneth B Fallon⁷⁵, David W Fardo⁷⁶, Martin R Farlow⁷⁷, Victoria Fernandez-Hernandez³⁶, Steven Ferris⁷⁸, Robert P Friedland⁷⁹, Tatiana M Foroud⁷³, Matthew P Frosch⁸⁰, Brian Fulton-Howard⁸¹, Douglas R Galasko⁴⁵, Adriana Gamboa^{82

83}, Marla Gearing^{84

85}, Daniel H Geschwind⁵⁶, Bernardino Ghetti⁸⁶, John R Gilbert^{1

2}, Rodney C P Go⁷⁵, Alison M Goate⁴⁸, Thomas J Grabowski^{39

87}, Neill R Graff-Radford^{21

71}, Robert C Green⁸⁸, John H Growdon⁸⁹, Hakon Hakonarson^{90

91}, James Hall²², Ronald L Hamilton⁹², Oscar Harari⁶³, John Hardy^{93

94}, Lindy E Harrell⁹⁵, Elizabeth Head⁹⁶, Victor W Henderson^{97

98}, Michelle Hernandez⁶⁸, Timothy Hohman^{99

100}, Lawrence S Honig⁴, Ryan M Huebinger¹⁰¹, Matthew J Huentelman¹⁰², Christine M Hulette¹⁰³, Bradley T Hyman⁸⁹, Linda S Hynan^{15

104

105}, Laura Ibanez^{106

107}, Gail P Jarvik^{108

109}, Suman Jayadev³⁹, Lee-Way Jin¹¹⁰, Kim Johnson⁶⁸, Leigh Johnson⁸², M Ilyas Kamboh^{111

112

113}, Anna M Karydas⁴⁴, Mindy J Katz¹¹⁴, John S Kauwe^{115

116}, Jeffrey A Kaye^{117

118}, C Dirk Keene¹¹⁹, Aisha Khaleeq⁶⁴, Masataka Kikuchi¹³, Ronald Kim⁹⁶, Janice Knebl⁸², Neil W Kowall^{10

120}, Joel H Kramer¹²¹, Walter A Kukull¹²², Frank M LaFerla¹²³, James J Lah¹²⁴, Eric B Larson¹²⁵, Alan Lerner³, James B Leverenz¹¹, Allan I Levey¹²⁴, Andrew P Lieberman¹²⁶, Richard B Lipton¹¹⁴, Mark Logue^{7

127

128}, Oscar L Lopez³⁴, Kathryn L Lunetta¹², Constantine G Lyketsos¹²⁹, Douglas Mains^{82

83}, Flanagan E Margaret^{130

131}, Daniel C Marson⁹⁵, Eden Rr Martin^{1

2}, Frank Martiniuk¹³², Deborah C Mash¹³³, Eliezer Masliah^{45

134}, Paul Massman⁶⁴, Arjun Masurkar⁷⁸, Wayne C McCormick⁵⁹, Susan M McCurry¹³⁵, Andrew N McDavid⁵³, Stefan McDonough¹³⁶, Ann C McKee^{10

137}, Marsel Mesulam^{130

131}, Bruce L Miller¹³⁸, Carol A Miller¹³⁹, Joshua W Miller¹¹⁰, Thomas J Montine¹⁴⁰, Edwin S Monuki¹⁴¹, John C Morris^{51

107

142

143}, Shubhabrata Mukherjee⁵⁹, Amanda J Myers⁶¹, Trung Nguyen¹⁰⁴, Thomas Obisesan¹⁴⁴, Sid O'Bryant¹⁴⁵, John M Olichney¹⁴⁶, Marcia Ory¹⁴⁷, Raymond Palmer¹⁴⁸, Joseph E Parisi¹⁴⁹, Henry L Paulson^{18

20}, Valory Pavlik⁶⁴, David Paydarfar¹⁶, Victoria Perez⁶⁸, Elaine Peskind¹⁵⁰, Ronald C Petersen⁴², Helen Petrovitch¹⁵¹, Aimee Pierce⁶⁰, Marsha Polk¹⁴⁸, Wayne W Poon⁶⁹, Huntington Potter¹⁵², Liming Qu⁸, Mary Quiceno^{153

154}, Joseph F Quinn^{117

118}, Ashok Raj⁵², Murray Raskind¹⁵⁰, Eric M Reiman^{102

155

156

157}, Barry Reisberg^{78

158}, Joan S Reisch⁶⁵, John M Ringman¹⁵⁹, Erik D Roberson⁹⁵, Monica Rodriguear⁶⁴, Ekaterina Rogaeva¹⁶⁰, Howard J Rosen⁴⁴, Roger N Rosenberg¹⁰⁴, Donald R Royall¹⁶¹, Marwan Sabbagh¹⁶², A Dessa Sadovnick¹⁶³, Mark A Sager²⁷, Mary Sano⁵⁰, Andrew J Saykin^{73

164}, Julie A Schneider^{29

31

165}, Lon S Schneider^{57

166}, William W Seeley⁴⁴, Susan H Slifer², Scott Small^{4

5}, Amanda G Smith⁵², Janet P Smith⁶⁵, Joshua A Sonnen¹¹⁹, Salvatore Spina⁸⁶, Peter St George-Hyslop^{167

168}, Takiyah D Starks^{169

170}, Robert A Stern¹⁰, Alan B Stevens^{171

172

173}, Stephen M Strittmatter¹⁷⁴, David Sultzer¹⁷⁵, Russell H Swerdlow⁴⁷, Rudolph E Tanzi⁸⁹, Jeffrey L Tilson¹⁷⁶, John Q Trojanowski⁸, Juan C Troncoso¹⁷⁷, Magda Tsolaki¹⁷⁸, Debby W Tsuang^{38

150}, Vivianna M Van Deerlin⁸, Linda J van Eldik¹⁷⁹, Jeffery M Vance^{1

2}, Badri N Vardarajan⁴, Robert Vassar^{130

131}, Harry V Vinters^{159

180}, Jean-Paul Vonsattel⁴, Sandra Weintraub¹⁸¹, Kathleen A Welsh-Bohmer^{46

182}, Patrice L Whitehead², Ellen M Wijsman^{108

109

183}, Kirk C Wilhelmsen⁵⁵, Benjamin Williams¹⁸⁴, Jennifer Williamson⁴, Henrik Wilms⁶⁸, Thomas S Wingo¹²⁴, Thomas Wisniewski^{185

186}, Randall L Woltjer¹⁸⁴, Martin Woon³⁷, Clinton B Wright¹⁸⁷, Chuang-Kuo Wu⁶⁸, Steven G Younkin^{21

71}, Chang-En Yu⁵⁹, Lei Yu^{28

29

31}, Xiongwei Zhu¹⁸⁸, Brian W Kunkle^{1

2}, William S Bush^{3

9}, Akinori Miyashita¹³, Goldie S Byrd¹⁸⁹, Li-San Wang⁸, Lindsay A Farrer^{10

7

12

189

190}, Jonathan L Haines^{3

9}, Richard Mayeux⁴, Margaret A Pericak-Vance^{1

2}, Gerard D Schellenberg⁸, Gyungah R Jun^#^{7

12

189}, Christiane Reitz^#^{5

32

191}, Adam C Naj^#^{192

193}; Alzheimer’s Disease Genetics Consortium (ADGC)

Affiliations

¹ Dr. John T. Macdonald Foundation Department of Human Genetics, Miller School of Medicine, University of Miami, Miami, FL, USA.
² The John P. Hussman Institute for Human Genomics, University of Miami, Miami, FL, USA.
³ Department of Population and Quantitative Health Sciences, Case Western Reserve University, Cleveland, OH, USA.
⁴ Taub Institute for Research in Alzheimer's Disease and the Aging Brain, The Gertrude H. Sergievsky Center Department of Neurology, Columbia University, New York, NY, USA.
⁵ Department of Neurology, Columbia University, New York, NY, USA.
⁶ Department of Biostatistics, Epidemiology, and Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
⁷ Department of Medicine (Biomedical Genetics), Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
⁸ Penn Neurodegeneration Genomics Center, Department of Pathology and Laboratory Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
⁹ Cleveland Institute for Computational Biology, Case Western Reserve University, Cleveland, OH, USA.
¹⁰ Department of Neurology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹¹ Cleveland Clinic Lou Ruvo Center for Brain Health, Cleveland Clinic, Cleveland, OH, USA.
¹² Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA.
¹³ Molecular Genetics Division, Brain Research Institute, Niigata University, Niigata, Japan.
¹⁴ Sanders-Brown Center on Aging, Department of Epidemiology, College of Public Health, University of Kentucky, Lexington, KY, USA.
¹⁵ Department of Psychiatry, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁶ Department of Neurology, Dell Medical School, University of Texas at Austin, Austin, TX, USA.
¹⁷ Department of Neurology, Johns Hopkins University, Baltimore, MD, USA.
¹⁸ Department of Neurology, University of Michigan, Ann Arbor, MI, USA.
¹⁹ Geriatric Research, Education and Clinical Center (GRECC), VA Ann Arbor Healthcare System (VAAAHS), Ann Arbor, MI, USA.
²⁰ Michigan Alzheimer's Disease Center, University of Michigan, Ann Arbor, MI, USA.
²¹ Department of Neuroscience, Mayo Clinic, Jacksonville, FL, USA.
²² Department of Pharmacology and Neuroscience, University of North Texas Health Science Center, Fort Worth, TX, USA.
²³ Departments of Neurology, Radiology, and Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, IN, USA.
²⁴ Indiana Alzheimer's Disease Research Center, Indiana University School of Medicine, Indianapolis, IN, USA.
²⁵ Department of Psychiatry, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
²⁶ Geriatric Research, Education and Clinical Center (GRECC), University of Wisconsin, Madison, WI, USA.
²⁷ Department of Medicine, University of Wisconsin, Madison, WI, USA.
²⁸ Wisconsin Alzheimer's Disease Research Center, Madison, WI, USA.
²⁹ Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, USA.
³⁰ Department of Behavioral Sciences, Rush University Medical Center, Chicago, IL, USA.
³¹ Rush Alzheimer's Disease Center, Rush University Medical Center, Chicago, IL, USA.
³² Gertrude H. Sergievsky Center, Columbia University, New York, NY, USA.
³³ Civin Laboratory for Neuropathology, Banner Sun Health Research Institute, Phoenix, AZ, USA.
³⁴ Departments of Psychiatry, Neurology, and Psychology, University of Pittsburgh School of Medicine, Pennsylvania, WA, USA.
³⁵ National Alzheimer's Coordinating Center, University of Washington, Seattle, WA, USA.
³⁶ Department of Psychiatry and Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, MO, USA.
³⁷ Department of Psychiatry , University of Texas at Austin/Dell Medical School, Austin, TX, USA.
³⁸ VA Puget Sound Health Care System/GRECC, Seattle, WA, USA.
³⁹ Department of Neurology, University of Washington, Seattle, WA, USA.
⁴⁰ Department of Epidemiology, Harvard School of Public Health, Boston, MA, USA.
⁴¹ Department of Psychiatry, Massachusetts General Hospital/Harvard Medical School, Boston, MA, USA.
⁴² Department of Neurology, Mayo Clinic, Rochester, MN, USA.
⁴³ Swedish Medical Center, Seattle, WA, USA.
⁴⁴ Department of Neurology, University of California San Francisco, San Francisco, CA, USA.
⁴⁵ Department of Neurosciences, University of California San Diego, La Jolla, CA, USA.
⁴⁶ Department of Medicine, Duke University, Durham, NC, USA.
⁴⁷ University of Kansas Alzheimer's Disease Center, University of Kansas Medical Center, Kansas City, KS, USA.
⁴⁸ Department of Genetics and Genomic Sciences, Ronald M. Loeb Center for Alzheimer's Disease, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁴⁹ Department of Neuroscience, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁵⁰ Department of Psychiatry, Mount Sinai School of Medicine, New York, NY, USA.
⁵¹ Department of Pathology and Immunology, Washington University, St. Louis, MO, USA.
⁵² USF Health Byrd Alzheimer's Institute, University of South Florida, Tampa, FL, USA.
⁵³ Fred Hutchinson Cancer Research Center, Seattle, WA, USA.
⁵⁴ Mental Health and Behavioral Science Service, Bruce W. Carter VA Medical Center, Miami, FL, USA.
⁵⁵ Department of Genetics, University of North Carolina Chapel Hill, Chapel Hill, NC, USA.
⁵⁶ Neurogenetics Program, University of California Los Angeles, Los Angeles, CA, USA.
⁵⁷ University of California Los Angeles, University of Southern California, Los Angeles, CA, USA.
⁵⁸ Section of Gerontology and Geriatric Medicine Research, Wake Forest School of Medicine, Winston-Salem, NC, USA.
⁵⁹ Department of Medicine, University of Washington, Seattle, WA, USA.
⁶⁰ Department of Neurology, University of California Irvine, Irvine, CA, USA.
⁶¹ Department of Psychiatry and Behavioral Sciences, Miller School of Medicine, University of Miami, Miami, FL, USA.
⁶² NeuroGenomics and Informatics, Washington University, St Louis, MO, USA.
⁶³ Department of Psychiatry, Washington University in St. Louis, St Louis, MO, USA.
⁶⁴ Alzheimer's Disease and Memory Disorders Center, Baylor College of Medicine, Houston, TX, USA.
⁶⁵ Department of Population and Data Sciences, University of Texas Southwestern Medical Center, Dallas, TX, USA.
⁶⁶ Center for Translational and Computational Neuroimmunology, Department of Neurology, Columbia University Medical Center, New York, NY, USA.
⁶⁷ Department of Neurology, University of California Davis, Sacramento, CA, USA.
⁶⁸ Departments of Neurology, Pharmacology and Neuroscience, Texas Tech University Health Science Center, Lubbock, TX, USA.
⁶⁹ Institute for Memory Impairments and Neurological Disorders, University of California Irvine, Irvine, CA, USA.
⁷⁰ Wien Center for Alzheimer's Disease and Memory Disorders, Mount Sinai Medical Center, Miami Beach, FL, USA.
⁷¹ Department of Neurology, Mayo Clinic, Jacksonville, FL, USA.
⁷² Rush Institute for Healthy Aging, Department of Internal Medicine, Rush University Medical Center, Chicago, IL, USA.
⁷³ Department of Medical and Molecular Genetics, Indiana University, Indianapolis, IN, USA.
⁷⁴ Office of Strategy and Measurement, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁷⁵ Department of Pathology, University of Alabama at Birmingham, Birmingham, AL, USA.
⁷⁶ Sanders-Brown Center on Aging, Department of Biostatistics, College of Public Health, University of Kentucky, Lexington, KY, USA.
⁷⁷ Department of Neurology, Indiana University, Indianapolis, IN, USA.
⁷⁸ Department of Psychiatry, New York University, New York, NY, USA.
⁷⁹ Department of Neurology, University of Louisville School of Medicine, Lousiville, KY, USA.
⁸⁰ C.S. Kubik Laboratory for Neuropathology, Massachusetts General Hospital, Charlestown, MA, USA.
⁸¹ Department of Neuroscience, Ronald M. Loeb Center for Alzheimer's Disease, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁸² Department of Health Behavior and Health Systems, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁸³ Department of Health Management and Policy, School of Public Health, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁸⁴ Department of Pathology and Laboratory Medicine, Emory University, Atlanta, GA, USA.
⁸⁵ Emory Alzheimer's Disease Center, Emory University, Atlanta, GA, USA.
⁸⁶ Department of Pathology and Laboratory Medicine, Indiana University, Indianapolis, IN, USA.
⁸⁷ Department of Radiology, University of Washington, Seattle, WA, USA.
⁸⁸ Division of Genetics, Department of Medicine and Partners Center for Personalized Genetic Medicine, Division of Genetics, Department of Medicine and Partners Center for Personalized Genetic Medicine, Boston, MA, USA.
⁸⁹ Department of Neurology, Massachusetts General Hospital/Harvard Medical School, Boston, MA, USA.
⁹⁰ Center for Applied Genomics, Children's Hospital of Philadelphia, Philadelphia, PA, USA.
⁹¹ Division of Human Genetics, Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, UK.
⁹² Department of Pathology (Neuropathology), University of Pittsburgh, Pittsburgh, PA, UK.
⁹³ UCL Institute of Neurology, University College London, London, England, USA.
⁹⁴ Department of Molecular Neuroscience, UCL Institute of Neurology, University College London, London, England, USA.
⁹⁵ Department of Neurology, University of Alabama at Birmingham, Birmingham, AL, USA.
⁹⁶ Department of Pathology and Laboratory Medicine, University of California Irvine, Irvine, CA, USA.
⁹⁷ Department of Epidemiology and Population Health, Stanford University, Stanford, CA, USA.
⁹⁸ Department of Neurology and Neurological Sciences, Stanford University, Stanford, CA, USA.
⁹⁹ Vanderbilt Memory and Alzheimer's Center, Department of Neurology, Vanderbilt University Medical Center, Nashville, TN, USA.
¹⁰⁰ Vanderbilt Genetics Institute, Division of Genetic Medicine, Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, USA.
¹⁰¹ Department of Surgery, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰² Neurogenomics Division, Translational Genomics Research Institute, Phoenix, AZ, USA.
¹⁰³ Department of Pathology, Duke University, Durham, NC, USA.
¹⁰⁴ Department of Neurology, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰⁵ Department of Neurological Surgery, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰⁶ Department of Psychiatry, Washington University School of Medicine, St. Louis, MO, USA.
¹⁰⁷ Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, MO, USA.
¹⁰⁸ Department of Genome Sciences, University of Washington, Seattle, WA, USA.
¹⁰⁹ Department of Medicine (Medical Genetics), University of Washington, Seattle, WA, USA.
¹¹⁰ Department of Pathology and Laboratory Medicine, University of California Davis, Sacramento, CA, USA.
¹¹¹ Department of Psychiatry, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹² Department of Human Genetics, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹³ Alzheimer's Disease Research Center, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹⁴ Department of Neurology, Albert Einstein College of Medicine, New York, NY, USA.
¹¹⁵ Department of Neuroscience, Brigham Young University, Provo, UT, USA.
¹¹⁶ Department of Biology, yBrigham Young University, Provo, UT, USA.
¹¹⁷ Department of Neurology, Oregon Health and Science University, Portland, OR, USA.
¹¹⁸ Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA.
¹¹⁹ Department of Laboratory Medicine and Pathology, University of Washington, Seattle, WA, USA.
¹²⁰ Department of Pathology, Boston University, Boston, MA, USA.
¹²¹ Department of Neuropsychology, University of California San Francisco, San Francisco, CA, USA.
¹²² Department of Epidemiology, University of Washington, Seattle, Seattle, USA.
¹²³ Department of Neurobiology and Behavior, University of California Irvine, Irvine, CA, USA.
¹²⁴ Department of Neurology, Emory University, Atlanta, GA, USA.
¹²⁵ National Center for PTSD, Boston VA Healthcare System, Boston, MA, USA.
¹²⁶ Department of Pathology, University of Michigan, Ann Arbor, MI, USA.
¹²⁷ National Center for PTSD at Boston VA Healthcare System, Boston, MA, USA.
¹²⁸ Department of Psychiatry, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹²⁹ Department of Psychiatry, Johns Hopkins University, Baltimore, MD, USA.
¹³⁰ Department of Pathology, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹³¹ Cognitive Neurology and Alzheimer's Disease Center, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹³² Department of Medicine - Pulmonary, New York University, New York, NY, USA.
¹³³ Department of Neurology, Miller School of Medicine, University of Miami, Miami, FL, USA.
¹³⁴ Department of Pathology, University of California San Diego, La Jolla, CA, USA.
¹³⁵ School of Nursing Northwest Research Group on Aging, University of Washington, Seattle, WA, USA.
¹³⁶ Pfizer Worldwide Research and Development, New York, NY, USA.
¹³⁷ Department of Pathology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹³⁸ Weill Institute for Neurosciences, Memory and Aging Center, University of California San Francisco, San Francisco, CA, USA.
¹³⁹ Department of Pathology, University of Southern California, Los Angeles, CA, USA.
¹⁴⁰ Department of Pathology, Stanford University School of Medicine, Stanford, CA, USA.
¹⁴¹ Department of Pathology and Laboratory Medicine and Alzheimer's Disease Research Center, University of California Irvine, Irvine, CA, USA.
¹⁴² Department of Neurology, Washington University, St. Louis, MO, USA.
¹⁴³ Department of Psychiatry, Washington University School of Medicine, St. Louis Missouri, USA.
¹⁴⁴ Department of Research Regulatory Compliance, College of Medicine, Howard Unviersity, Washington, DC, USA.
¹⁴⁵ Institute for Translational Research, University of North Texas Health Science Center, Fort Worth,, TX, USA.
¹⁴⁶ Center for Mind and Brain and Department of Neurology, University of California Davis, Sacramento, CA, USA.
¹⁴⁷ Center for Population Health and Aging, Texas AandM University Health Science Center, Lubbock , TX, USA.
¹⁴⁸ Department of Family and Community Medicine, University of Texas Health Science Center San Antonio, San Antonio, TX, USA.
¹⁴⁹ Department of Laboratory Medicine and Pathology, Mayo Clinic, Rochester, MN, USA.
¹⁵⁰ Department of Psychiatry and Behavioral Sciences, University of Washington School of Medicine, Seattle, WA, USA.
¹⁵¹ Pacific Health Research & Education Institute, VA Pacific Islands Healthcare System, Honolulu, HI, USA.
¹⁵² Department of Neurology, University of Colorado School of Medicine, Aurora, CO, USA.
¹⁵³ Department of Internal Medicine and Geriatrics, University of North Texas Health Science Center, Fort Worth, TX, USA.
¹⁵⁴ Department of Medical Education, TCU/UNTHSC School of Medicine, Fort Worth, TX, USA.
¹⁵⁵ Arizona Alzheimer's Consortium, Phoenix, AZ, Canada.
¹⁵⁶ Banner Alzheimer's Institute, Phoenix, AZ, USA.
¹⁵⁷ Department of Psychiatry, University of Arizona, Phoenix, AZ, USA.
¹⁵⁸ Alzheimer's Disease Center, New York University, New York, NY, USA.
¹⁵⁹ Department of Neurology, University of California Los Angeles, Los Angeles, CA, USA.
¹⁶⁰ Tanz Centre for Research in Neurodegenerative Disease, University of Toronto, Toronto, ON, Canada.
¹⁶¹ Departments of Psychiatry, Medicine, Family and Community Medicine, and the Glenn Biggs Institute for Alzheimer's and Neurodegenerative Diseases, UT Health Science Center at San Antonio,, San Antonio, TX, Canada.
¹⁶² Department of Neurology, Barrow Neurological Institute St. Joseph's Hospital and Medical Center, Phoenix, AZ, USA.
¹⁶³ Department of Medical Genetics, University of British Columbia, Vancouver, Canada.
¹⁶⁴ Department of Radiology and Imaging Sciences, Indiana University, Indianapolis, IN, USA.
¹⁶⁵ Department of Pathology (Neuropathology), Rush University Medical Center, Chicago, IL, USA.
¹⁶⁶ Department of Psychiatry, University of Southern California, Los Angeles, CA, USA.
¹⁶⁷ Cambridge Institute for Medical Research, University of Cambridge, Cambridge, England, USA.
¹⁶⁸ Faculty of Medicine, Department of Medicine (Neurology), University of Toronto, Toronto, ON, USA.
¹⁶⁹ Maya Angelou Center for Health Equity, Wake Forest School of Medicine, Winston-Salem, NC, USA.
¹⁷⁰ Center for Outreach in Alzheimer's, Aging and Community Health at North Carolina A&T State University, Greensboro, NC, USA.
¹⁷¹ Center for Applied Health Research, Baylor Scott & White Health, Temple, TX, USA.
¹⁷² Center for Population Health and Aging, Texas A&M University Health Science Center, Lubbock, TX, USA.
¹⁷³ College of Medicine, Texas A&M University Health Science Center, College Station, TX, USA.
¹⁷⁴ Program in Cellular Neuroscience, Neurodegeneration and Repair, Yale University School of Medicine, New Haven, CT, USA.
¹⁷⁵ Department of Psychiatry and Human Behavior, University of California Irvine, Irvine, CA, USA.
¹⁷⁶ Renaissance Computing Institute, University of North Carolina Chapel Hill, Chapel Hill, NC, USA.
¹⁷⁷ Department of Pathology, Johns Hopkins University, Baltimore, MD, USA.
¹⁷⁸ Department of Neurology, Aristotle University of Thessaloniki, Thessaloniki, Macedonia, USA.
¹⁷⁹ Sanders-Brown Center on Aging, Department of Neuroscience, College of Medicine, University of Kentucky, Kentucky, OH, USA.
¹⁸⁰ Department of Pathology and Laboratory Medicine, University of California Los Angeles, Los Angeles, CA, USA.
¹⁸¹ Department of Psychiatry and Behavioral Sciences, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹⁸² Department of Psychiatry and Behavioral Sciences, Duke University, Durham, NC, USA.
¹⁸³ Department of Biostatistics, University of Washington, Seattle, WA, USA.
¹⁸⁴ Department of Pathology, Oregon Health and Science University, Portland, OR, USA.
¹⁸⁵ Department of Psychiatry, New York University Grossman School of Medicine, New York, NY, USA.
¹⁸⁶ Center for Cognitive Neurology and Departments of Neurology and Pathology, New York University Grossman School of Medicine, New York, USA.
¹⁸⁷ Evelyn F. McKnight Brain Institute, Department of Neurology, Miller School of Medicine, University of Miami, Miami, FL, USA.
¹⁸⁸ Social Sciences & Health Policy, Wake Forest School of Medicine, Winston-Salem, NC, USA.
¹⁸⁹ Department of Ophthalmology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹⁹⁰ Department of Epidemiology, Boston University School of Public Health, Boston, MA, USA.
¹⁹¹ Department of Epidemiology, Columbia University, New York, NY, USA.
¹⁹² Department of Biostatistics, Epidemiology, and Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA. adamnaj@pennmedicine.upenn.edu.
¹⁹³ Penn Neurodegeneration Genomics Center, Department of Pathology and Laboratory Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA. adamnaj@pennmedicine.upenn.edu.

^# Contributed equally.

PMID: 40676597
PMCID: PMC12273372
DOI: 10.1186/s13059-025-03564-z

Meta-Analysis

Multi-ancestry genome-wide meta-analysis of 56,241 individuals identifies known and novel cross-population and ancestry-specific associations as novel risk loci for Alzheimer's disease

Farid Rajabli et al. Genome Biol. 2025.

. 2025 Jul 17;26(1):210.

doi: 10.1186/s13059-025-03564-z.

Authors

Affiliations

¹ Dr. John T. Macdonald Foundation Department of Human Genetics, Miller School of Medicine, University of Miami, Miami, FL, USA.
² The John P. Hussman Institute for Human Genomics, University of Miami, Miami, FL, USA.
³ Department of Population and Quantitative Health Sciences, Case Western Reserve University, Cleveland, OH, USA.
⁴ Taub Institute for Research in Alzheimer's Disease and the Aging Brain, The Gertrude H. Sergievsky Center Department of Neurology, Columbia University, New York, NY, USA.
⁵ Department of Neurology, Columbia University, New York, NY, USA.
⁶ Department of Biostatistics, Epidemiology, and Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
⁷ Department of Medicine (Biomedical Genetics), Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
⁸ Penn Neurodegeneration Genomics Center, Department of Pathology and Laboratory Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
⁹ Cleveland Institute for Computational Biology, Case Western Reserve University, Cleveland, OH, USA.
¹⁰ Department of Neurology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹¹ Cleveland Clinic Lou Ruvo Center for Brain Health, Cleveland Clinic, Cleveland, OH, USA.
¹² Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA.
¹³ Molecular Genetics Division, Brain Research Institute, Niigata University, Niigata, Japan.
¹⁴ Sanders-Brown Center on Aging, Department of Epidemiology, College of Public Health, University of Kentucky, Lexington, KY, USA.
¹⁵ Department of Psychiatry, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁶ Department of Neurology, Dell Medical School, University of Texas at Austin, Austin, TX, USA.
¹⁷ Department of Neurology, Johns Hopkins University, Baltimore, MD, USA.
¹⁸ Department of Neurology, University of Michigan, Ann Arbor, MI, USA.
¹⁹ Geriatric Research, Education and Clinical Center (GRECC), VA Ann Arbor Healthcare System (VAAAHS), Ann Arbor, MI, USA.
²⁰ Michigan Alzheimer's Disease Center, University of Michigan, Ann Arbor, MI, USA.
²¹ Department of Neuroscience, Mayo Clinic, Jacksonville, FL, USA.
²² Department of Pharmacology and Neuroscience, University of North Texas Health Science Center, Fort Worth, TX, USA.
²³ Departments of Neurology, Radiology, and Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, IN, USA.
²⁴ Indiana Alzheimer's Disease Research Center, Indiana University School of Medicine, Indianapolis, IN, USA.
²⁵ Department of Psychiatry, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.
²⁶ Geriatric Research, Education and Clinical Center (GRECC), University of Wisconsin, Madison, WI, USA.
²⁷ Department of Medicine, University of Wisconsin, Madison, WI, USA.
²⁸ Wisconsin Alzheimer's Disease Research Center, Madison, WI, USA.
²⁹ Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, USA.
³⁰ Department of Behavioral Sciences, Rush University Medical Center, Chicago, IL, USA.
³¹ Rush Alzheimer's Disease Center, Rush University Medical Center, Chicago, IL, USA.
³² Gertrude H. Sergievsky Center, Columbia University, New York, NY, USA.
³³ Civin Laboratory for Neuropathology, Banner Sun Health Research Institute, Phoenix, AZ, USA.
³⁴ Departments of Psychiatry, Neurology, and Psychology, University of Pittsburgh School of Medicine, Pennsylvania, WA, USA.
³⁵ National Alzheimer's Coordinating Center, University of Washington, Seattle, WA, USA.
³⁶ Department of Psychiatry and Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, MO, USA.
³⁷ Department of Psychiatry , University of Texas at Austin/Dell Medical School, Austin, TX, USA.
³⁸ VA Puget Sound Health Care System/GRECC, Seattle, WA, USA.
³⁹ Department of Neurology, University of Washington, Seattle, WA, USA.
⁴⁰ Department of Epidemiology, Harvard School of Public Health, Boston, MA, USA.
⁴¹ Department of Psychiatry, Massachusetts General Hospital/Harvard Medical School, Boston, MA, USA.
⁴² Department of Neurology, Mayo Clinic, Rochester, MN, USA.
⁴³ Swedish Medical Center, Seattle, WA, USA.
⁴⁴ Department of Neurology, University of California San Francisco, San Francisco, CA, USA.
⁴⁵ Department of Neurosciences, University of California San Diego, La Jolla, CA, USA.
⁴⁶ Department of Medicine, Duke University, Durham, NC, USA.
⁴⁷ University of Kansas Alzheimer's Disease Center, University of Kansas Medical Center, Kansas City, KS, USA.
⁴⁸ Department of Genetics and Genomic Sciences, Ronald M. Loeb Center for Alzheimer's Disease, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁴⁹ Department of Neuroscience, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁵⁰ Department of Psychiatry, Mount Sinai School of Medicine, New York, NY, USA.
⁵¹ Department of Pathology and Immunology, Washington University, St. Louis, MO, USA.
⁵² USF Health Byrd Alzheimer's Institute, University of South Florida, Tampa, FL, USA.
⁵³ Fred Hutchinson Cancer Research Center, Seattle, WA, USA.
⁵⁴ Mental Health and Behavioral Science Service, Bruce W. Carter VA Medical Center, Miami, FL, USA.
⁵⁵ Department of Genetics, University of North Carolina Chapel Hill, Chapel Hill, NC, USA.
⁵⁶ Neurogenetics Program, University of California Los Angeles, Los Angeles, CA, USA.
⁵⁷ University of California Los Angeles, University of Southern California, Los Angeles, CA, USA.
⁵⁸ Section of Gerontology and Geriatric Medicine Research, Wake Forest School of Medicine, Winston-Salem, NC, USA.
⁵⁹ Department of Medicine, University of Washington, Seattle, WA, USA.
⁶⁰ Department of Neurology, University of California Irvine, Irvine, CA, USA.
⁶¹ Department of Psychiatry and Behavioral Sciences, Miller School of Medicine, University of Miami, Miami, FL, USA.
⁶² NeuroGenomics and Informatics, Washington University, St Louis, MO, USA.
⁶³ Department of Psychiatry, Washington University in St. Louis, St Louis, MO, USA.
⁶⁴ Alzheimer's Disease and Memory Disorders Center, Baylor College of Medicine, Houston, TX, USA.
⁶⁵ Department of Population and Data Sciences, University of Texas Southwestern Medical Center, Dallas, TX, USA.
⁶⁶ Center for Translational and Computational Neuroimmunology, Department of Neurology, Columbia University Medical Center, New York, NY, USA.
⁶⁷ Department of Neurology, University of California Davis, Sacramento, CA, USA.
⁶⁸ Departments of Neurology, Pharmacology and Neuroscience, Texas Tech University Health Science Center, Lubbock, TX, USA.
⁶⁹ Institute for Memory Impairments and Neurological Disorders, University of California Irvine, Irvine, CA, USA.
⁷⁰ Wien Center for Alzheimer's Disease and Memory Disorders, Mount Sinai Medical Center, Miami Beach, FL, USA.
⁷¹ Department of Neurology, Mayo Clinic, Jacksonville, FL, USA.
⁷² Rush Institute for Healthy Aging, Department of Internal Medicine, Rush University Medical Center, Chicago, IL, USA.
⁷³ Department of Medical and Molecular Genetics, Indiana University, Indianapolis, IN, USA.
⁷⁴ Office of Strategy and Measurement, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁷⁵ Department of Pathology, University of Alabama at Birmingham, Birmingham, AL, USA.
⁷⁶ Sanders-Brown Center on Aging, Department of Biostatistics, College of Public Health, University of Kentucky, Lexington, KY, USA.
⁷⁷ Department of Neurology, Indiana University, Indianapolis, IN, USA.
⁷⁸ Department of Psychiatry, New York University, New York, NY, USA.
⁷⁹ Department of Neurology, University of Louisville School of Medicine, Lousiville, KY, USA.
⁸⁰ C.S. Kubik Laboratory for Neuropathology, Massachusetts General Hospital, Charlestown, MA, USA.
⁸¹ Department of Neuroscience, Ronald M. Loeb Center for Alzheimer's Disease, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
⁸² Department of Health Behavior and Health Systems, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁸³ Department of Health Management and Policy, School of Public Health, University of North Texas Health Science Center, Fort Worth, TX, USA.
⁸⁴ Department of Pathology and Laboratory Medicine, Emory University, Atlanta, GA, USA.
⁸⁵ Emory Alzheimer's Disease Center, Emory University, Atlanta, GA, USA.
⁸⁶ Department of Pathology and Laboratory Medicine, Indiana University, Indianapolis, IN, USA.
⁸⁷ Department of Radiology, University of Washington, Seattle, WA, USA.
⁸⁸ Division of Genetics, Department of Medicine and Partners Center for Personalized Genetic Medicine, Division of Genetics, Department of Medicine and Partners Center for Personalized Genetic Medicine, Boston, MA, USA.
⁸⁹ Department of Neurology, Massachusetts General Hospital/Harvard Medical School, Boston, MA, USA.
⁹⁰ Center for Applied Genomics, Children's Hospital of Philadelphia, Philadelphia, PA, USA.
⁹¹ Division of Human Genetics, Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, UK.
⁹² Department of Pathology (Neuropathology), University of Pittsburgh, Pittsburgh, PA, UK.
⁹³ UCL Institute of Neurology, University College London, London, England, USA.
⁹⁴ Department of Molecular Neuroscience, UCL Institute of Neurology, University College London, London, England, USA.
⁹⁵ Department of Neurology, University of Alabama at Birmingham, Birmingham, AL, USA.
⁹⁶ Department of Pathology and Laboratory Medicine, University of California Irvine, Irvine, CA, USA.
⁹⁷ Department of Epidemiology and Population Health, Stanford University, Stanford, CA, USA.
⁹⁸ Department of Neurology and Neurological Sciences, Stanford University, Stanford, CA, USA.
⁹⁹ Vanderbilt Memory and Alzheimer's Center, Department of Neurology, Vanderbilt University Medical Center, Nashville, TN, USA.
¹⁰⁰ Vanderbilt Genetics Institute, Division of Genetic Medicine, Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, USA.
¹⁰¹ Department of Surgery, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰² Neurogenomics Division, Translational Genomics Research Institute, Phoenix, AZ, USA.
¹⁰³ Department of Pathology, Duke University, Durham, NC, USA.
¹⁰⁴ Department of Neurology, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰⁵ Department of Neurological Surgery, University of Texas Southwestern Medical Center, Dallas, TX, USA.
¹⁰⁶ Department of Psychiatry, Washington University School of Medicine, St. Louis, MO, USA.
¹⁰⁷ Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, MO, USA.
¹⁰⁸ Department of Genome Sciences, University of Washington, Seattle, WA, USA.
¹⁰⁹ Department of Medicine (Medical Genetics), University of Washington, Seattle, WA, USA.
¹¹⁰ Department of Pathology and Laboratory Medicine, University of California Davis, Sacramento, CA, USA.
¹¹¹ Department of Psychiatry, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹² Department of Human Genetics, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹³ Alzheimer's Disease Research Center, University of Pittsburgh, Pittsburgh, PA, USA.
¹¹⁴ Department of Neurology, Albert Einstein College of Medicine, New York, NY, USA.
¹¹⁵ Department of Neuroscience, Brigham Young University, Provo, UT, USA.
¹¹⁶ Department of Biology, yBrigham Young University, Provo, UT, USA.
¹¹⁷ Department of Neurology, Oregon Health and Science University, Portland, OR, USA.
¹¹⁸ Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA.
¹¹⁹ Department of Laboratory Medicine and Pathology, University of Washington, Seattle, WA, USA.
¹²⁰ Department of Pathology, Boston University, Boston, MA, USA.
¹²¹ Department of Neuropsychology, University of California San Francisco, San Francisco, CA, USA.
¹²² Department of Epidemiology, University of Washington, Seattle, Seattle, USA.
¹²³ Department of Neurobiology and Behavior, University of California Irvine, Irvine, CA, USA.
¹²⁴ Department of Neurology, Emory University, Atlanta, GA, USA.
¹²⁵ National Center for PTSD, Boston VA Healthcare System, Boston, MA, USA.
¹²⁶ Department of Pathology, University of Michigan, Ann Arbor, MI, USA.
¹²⁷ National Center for PTSD at Boston VA Healthcare System, Boston, MA, USA.
¹²⁸ Department of Psychiatry, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹²⁹ Department of Psychiatry, Johns Hopkins University, Baltimore, MD, USA.
¹³⁰ Department of Pathology, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹³¹ Cognitive Neurology and Alzheimer's Disease Center, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹³² Department of Medicine - Pulmonary, New York University, New York, NY, USA.
¹³³ Department of Neurology, Miller School of Medicine, University of Miami, Miami, FL, USA.
¹³⁴ Department of Pathology, University of California San Diego, La Jolla, CA, USA.
¹³⁵ School of Nursing Northwest Research Group on Aging, University of Washington, Seattle, WA, USA.
¹³⁶ Pfizer Worldwide Research and Development, New York, NY, USA.
¹³⁷ Department of Pathology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹³⁸ Weill Institute for Neurosciences, Memory and Aging Center, University of California San Francisco, San Francisco, CA, USA.
¹³⁹ Department of Pathology, University of Southern California, Los Angeles, CA, USA.
¹⁴⁰ Department of Pathology, Stanford University School of Medicine, Stanford, CA, USA.
¹⁴¹ Department of Pathology and Laboratory Medicine and Alzheimer's Disease Research Center, University of California Irvine, Irvine, CA, USA.
¹⁴² Department of Neurology, Washington University, St. Louis, MO, USA.
¹⁴³ Department of Psychiatry, Washington University School of Medicine, St. Louis Missouri, USA.
¹⁴⁴ Department of Research Regulatory Compliance, College of Medicine, Howard Unviersity, Washington, DC, USA.
¹⁴⁵ Institute for Translational Research, University of North Texas Health Science Center, Fort Worth,, TX, USA.
¹⁴⁶ Center for Mind and Brain and Department of Neurology, University of California Davis, Sacramento, CA, USA.
¹⁴⁷ Center for Population Health and Aging, Texas AandM University Health Science Center, Lubbock , TX, USA.
¹⁴⁸ Department of Family and Community Medicine, University of Texas Health Science Center San Antonio, San Antonio, TX, USA.
¹⁴⁹ Department of Laboratory Medicine and Pathology, Mayo Clinic, Rochester, MN, USA.
¹⁵⁰ Department of Psychiatry and Behavioral Sciences, University of Washington School of Medicine, Seattle, WA, USA.
¹⁵¹ Pacific Health Research & Education Institute, VA Pacific Islands Healthcare System, Honolulu, HI, USA.
¹⁵² Department of Neurology, University of Colorado School of Medicine, Aurora, CO, USA.
¹⁵³ Department of Internal Medicine and Geriatrics, University of North Texas Health Science Center, Fort Worth, TX, USA.
¹⁵⁴ Department of Medical Education, TCU/UNTHSC School of Medicine, Fort Worth, TX, USA.
¹⁵⁵ Arizona Alzheimer's Consortium, Phoenix, AZ, Canada.
¹⁵⁶ Banner Alzheimer's Institute, Phoenix, AZ, USA.
¹⁵⁷ Department of Psychiatry, University of Arizona, Phoenix, AZ, USA.
¹⁵⁸ Alzheimer's Disease Center, New York University, New York, NY, USA.
¹⁵⁹ Department of Neurology, University of California Los Angeles, Los Angeles, CA, USA.
¹⁶⁰ Tanz Centre for Research in Neurodegenerative Disease, University of Toronto, Toronto, ON, Canada.
¹⁶¹ Departments of Psychiatry, Medicine, Family and Community Medicine, and the Glenn Biggs Institute for Alzheimer's and Neurodegenerative Diseases, UT Health Science Center at San Antonio,, San Antonio, TX, Canada.
¹⁶² Department of Neurology, Barrow Neurological Institute St. Joseph's Hospital and Medical Center, Phoenix, AZ, USA.
¹⁶³ Department of Medical Genetics, University of British Columbia, Vancouver, Canada.
¹⁶⁴ Department of Radiology and Imaging Sciences, Indiana University, Indianapolis, IN, USA.
¹⁶⁵ Department of Pathology (Neuropathology), Rush University Medical Center, Chicago, IL, USA.
¹⁶⁶ Department of Psychiatry, University of Southern California, Los Angeles, CA, USA.
¹⁶⁷ Cambridge Institute for Medical Research, University of Cambridge, Cambridge, England, USA.
¹⁶⁸ Faculty of Medicine, Department of Medicine (Neurology), University of Toronto, Toronto, ON, USA.
¹⁶⁹ Maya Angelou Center for Health Equity, Wake Forest School of Medicine, Winston-Salem, NC, USA.
¹⁷⁰ Center for Outreach in Alzheimer's, Aging and Community Health at North Carolina A&T State University, Greensboro, NC, USA.
¹⁷¹ Center for Applied Health Research, Baylor Scott & White Health, Temple, TX, USA.
¹⁷² Center for Population Health and Aging, Texas A&M University Health Science Center, Lubbock, TX, USA.
¹⁷³ College of Medicine, Texas A&M University Health Science Center, College Station, TX, USA.
¹⁷⁴ Program in Cellular Neuroscience, Neurodegeneration and Repair, Yale University School of Medicine, New Haven, CT, USA.
¹⁷⁵ Department of Psychiatry and Human Behavior, University of California Irvine, Irvine, CA, USA.
¹⁷⁶ Renaissance Computing Institute, University of North Carolina Chapel Hill, Chapel Hill, NC, USA.
¹⁷⁷ Department of Pathology, Johns Hopkins University, Baltimore, MD, USA.
¹⁷⁸ Department of Neurology, Aristotle University of Thessaloniki, Thessaloniki, Macedonia, USA.
¹⁷⁹ Sanders-Brown Center on Aging, Department of Neuroscience, College of Medicine, University of Kentucky, Kentucky, OH, USA.
¹⁸⁰ Department of Pathology and Laboratory Medicine, University of California Los Angeles, Los Angeles, CA, USA.
¹⁸¹ Department of Psychiatry and Behavioral Sciences, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
¹⁸² Department of Psychiatry and Behavioral Sciences, Duke University, Durham, NC, USA.
¹⁸³ Department of Biostatistics, University of Washington, Seattle, WA, USA.
¹⁸⁴ Department of Pathology, Oregon Health and Science University, Portland, OR, USA.
¹⁸⁵ Department of Psychiatry, New York University Grossman School of Medicine, New York, NY, USA.
¹⁸⁶ Center for Cognitive Neurology and Departments of Neurology and Pathology, New York University Grossman School of Medicine, New York, USA.
¹⁸⁷ Evelyn F. McKnight Brain Institute, Department of Neurology, Miller School of Medicine, University of Miami, Miami, FL, USA.
¹⁸⁸ Social Sciences & Health Policy, Wake Forest School of Medicine, Winston-Salem, NC, USA.
¹⁸⁹ Department of Ophthalmology, Boston University Chobanian & Avedisian School of Medicine, Boston, MA, USA.
¹⁹⁰ Department of Epidemiology, Boston University School of Public Health, Boston, MA, USA.
¹⁹¹ Department of Epidemiology, Columbia University, New York, NY, USA.
¹⁹² Department of Biostatistics, Epidemiology, and Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA. adamnaj@pennmedicine.upenn.edu.
¹⁹³ Penn Neurodegeneration Genomics Center, Department of Pathology and Laboratory Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA. adamnaj@pennmedicine.upenn.edu.

^# Contributed equally.

PMID: 40676597
PMCID: PMC12273372
DOI: 10.1186/s13059-025-03564-z

Abstract

Background: Limited ancestral diversity has impaired our ability to detect risk variants more prevalent in ancestry groups of predominantly non-European ancestral background in genome-wide association studies (GWAS). We construct and analyze a multi-ancestry GWAS dataset in the Alzheimer's Disease Genetics Consortium (ADGC) to test for novel shared and population-specific late-onset Alzheimer's disease (LOAD) susceptibility loci and evaluate underlying genetic architecture in 37,382 non-Hispanic White (NHW), 6728 African American, 8899 Hispanic (HIS), and 3232 East Asian individuals, performing within ancestry fixed-effects meta-analysis followed by a cross-ancestry random-effects meta-analysis.

Results: We identify 13 loci with cross-population associations including known loci at/near CR1, BIN1, TREM2, CD2AP, PTK2B, CLU, SHARPIN, MS4A6A, PICALM, ABCA7, APOE, and two novel loci not previously reported at 11p12 (LRRC4C) and 12q24.13 (LHX5-AS1). We additionally identify three population-specific loci with genome-wide significance at/near PTPRK and GRB14 in HIS and KIAA0825 in NHW. Pathway analysis implicates multiple amyloid regulation pathways and the classical complement pathway. Genes at/near our novel loci have known roles in neuronal development (LRRC4C, LHX5-AS1, and PTPRK) and insulin receptor activity regulation (GRB14).

Conclusions: Using cross-population GWAS meta-analyses, we identify novel LOAD susceptibility loci in/near LRRC4C and LHX5-AS1, both with known roles in neuronal development, as well as several novel population-unique loci. Reflecting the power of diverse ancestry in GWAS, we detect the SHARPIN locus with only 13.7% of the sample size of the NHW GWAS study (n = 409,589) in which this locus was first observed. Continued expansion into larger multi-ancestry studies will provide even more power for further elucidating the genomics of late-onset Alzheimer's disease.

Keywords: Alzheimer disease; GWAS meta-analysis,; Genome-wide association study (GWAS); Multi-ancestry; Population-specific.

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Conflict of interest statement

Declarations. Ethics approval and consent to participate: For our IRB to compile the de-identified data and conduct analyses, the ADGC protocol is reviewed and approved by the University of Pennsylvania IRB #8 (Federalwide Assurance #00004028). Consent for publication: Not applicable. Competing interests: J.A.P. has received compensation for serving as a section editor for Springer Nature and a grant reviewer with the Department of Defense and Research Grants Council of Hong Kong. M.S.A. is an advisor to Eli Lilly. L.G.A. receives compensation as a consultant for Biogen, Two Labs, IQVIA, NIH, Florida Department of Health, NIH Biobank, Eli Lilly, GE Healthcare, and Eisai; has received compensation for lectures, etc. from AAN, MillerMed, AiSM, and Health and Hospitality; and has received travel and meeting support from the Alzheimer’s Association; she also participates on Data Safety Monitoring or Advisory boards for IQVIA, NIA R01 AG061111, the UAB Nathan Shock Center, and the New Mexico Exploratory ADRC; she has received compensation for leadership roles in the Medical Science Council Alzheimer Association Greater IN Chapter, the Alzheimer Association Science Program Committee, and the FDA PCNS Advisory Committee; she also has stock or stock options Cassava Neurosciences and Golden Seeds; and has received materials support from AVID Pharmaceuticals, Life Molecular Imaging, and Roche Diagnostics. S.E.A. has received honoraria and/or travel expenses for lectures from Abbvie, Eisai, and Biogen and has served on scientific advisory boards of Corte, has received consulting fees from Athira, Cassava, Cognito Therapeutics, EIP Pharma and Orthogonal Neuroscience, and has received research grant support from NIH, Alzheimer’s Association, Alzheimer’s Drug Discovery Foundation, Abbvie, Amylyx, EIP Pharma, Merck, Janssen/Johnson & Johnson, Novartis, and vTv. S.Asthana reported receiving grants from National Institute on Aging/National Institutes of Health, Genentech, Merck, Toyoma Chemical, and Lundbeck outside the submitted work. L.L.B. has served as deputy editor for Alzheimer’s and Dementia for the Alzheimer’s Association. D.A.B. is a consultant for Biogen, Inc. B.F.B. has received institutional support from LBDA; is a member of the Scientific Advisory Boards of the Tau Consortium (funded by the Rainwater Charitable Foundation), AFTD, LBDA, and GE Healthcare; is a member of the Data Safety Monitoring Board of trial involving mesenchymal stem cells in MSA. J.D.B has received honoraria from serving on the Scientific Advisory Board and Speaker’s Bureau of Biogen, Celgene, EMD Serono, Genentech and Novartis; has received research support from AbbVie, Alexion, Alkermes, Biogen, Celgene, Sanofi Genzyme, Genentech, Novartis and TG Therapeutics. A.L.B. has received financial support from NIH, the Association for Frontotemporal Degeneration, the Bluefield Project, the Rainwater Charitable Foundation, Regeneron, Eisai and Biogen; and has served as a paid consultant for AGTC, Alector, Amylyx, AviadoBio, Arkuda, Arrowhead, Arvinas, Eli Lilly, Genentech, LifeEdit, Merck, Modalis, Oligomerix, Oscotec, Transposon and Wave. J.M.B. is compensated as a consultant for Stage 2 Innovations; and has received honoraria and travel support for speaking from Astra-Zeneca. J.D.Buxbaum is a consultant to BridgeBio and to Rumi; holds a patent for IGF- 1 in Phelan-McDermid syndrome; holds an honorary professorship from Aarhus University Denmark; receives research support from Takeda and Oryzon; and is a journal editor for Springer Nature. C.Cao has a patent pending for melatonin-insulin-THC (MIT) treatment; and serves as a scientific consultant for MegaNano Biotech, Inc. C.M.C. has received grants from the National Institutes of Health, Eisai, Eli Lilly, Veterans Affairs; has received nonfinancial support from Amarin; has received data safety monitoring board/travel/advisory board honoraria from Alzheimer's Association, National Institutes of Health, and American Fed Aging Res Beeson Program. J.C. is currently employed as a senior scientist at Takeda Pharmaceuticals, Inc.; the company did not influence the study design, analyses, or interpretation of the results presented in this manuscript. C.Cruchaga has received research support from GSK and Eisai, is a member of the advisory boards of Vivid Genomics and Circular Genomics, and owns stocks. D.W.D. is an editorial board member for Acta Neuropathologica, Brain, Brain Pathology, Neuropathology and Applied Neurobiology, Annals of Neurology, Neuropathology, and is an Editor for the International Journal of Clinical and Experimental Pathology and for the American Journal of Neurodegenerative Disease; and receives support Mangurian Foundation and the Rainwater Charitable Foundation. R.S.D. is an employee of F. Hoffman-La Roche, Ltd. and Genentech, Inc.; and owns or has stock options in F. Hoffmann-La Roche, Ltd. N.E.-T. receives research support from the NIH; is a member of multiple Scientific Advisory Boards including the Framingham Heart Study Executive Committee, Cytox, and the NIH TREAT-AD Consortium External Advisory Board Member; has patents pending for Human Monoclonal Antibodies Against Amyloid Beta Protein and Their Use as Therapeutic Agents Application, and RNAi against targets in Progressive Supranuclear palsy; is an Editorial Board Member for the American Journal of Neurodegenerative Disease and Alzheimer's & Dementia; and receives research support from Florida Health Ed and Ethel Moore Alzheimer's Disease Research Program and an Alzheimer's Association Zenith Award. T.M.F. has received honoraria and travel support from the External Advisory Boards for Alzheimer Disease Research Centers that might also be a site for the LEADS study. D.R.G. serves on Data Safety Monitoring Boards for Cognition Therapeutics and Proclara Biosciences; and is an Editor of Alzheimer’s Research & Therapy. B.G. has consulted for Piramal Imaging. A.M.G is a member of the Scientific Advisory Boards/Scientific Research Boards for Genentech, Muna Therapeutics, and Denali Therapeutics. N.R.G.-.R. has received royalties for an article in UpToDate; and has received research support for multi-center studies at Eli Lilly & Company, Biogen, and AbbVie. R.C.G. has received compensation as an advisor for AIA, Grail, Humanity, Kneed Media, Plumcare, UnitedHealth, Verily, VibrentHealth, Wamberg; and is co-founder of Genome Medical, Inc. J.Hardy is supported by the UK Dementia Research Institute, which receives its funding from DRI, Ltd., funded by the UK Medical Research Council, Alzheimer's Society, and Alzheimer's Research UK; and is also supported by the MRC, Wellcome Trust, the Dolby Family Fund, and the National Institute for Health Research University College London Hospitals Biomedical Research Centre. T.H. is a member of a scientific advisory board for Vivid Genomics. L.S.H. is the Web Editor for JAMA Neurology. B.T.H. has a family member who works at Novartis, and owns stock in Novartis; and serves on the Scientific Advisory Board of Dewpoint and owns stock; and serves on a Scientific Advisory Board or is a consultant for AbbVie, Aprinoia Therapeutics, Arvinas, Avrobio, Axial, Biogen, BMS, Cure Alz Fund, Cell Signaling, Eisai, Genentech, Ionis, Latus, Novartis, Sangamo, Sanofi, Seer, Takeda, the US Dept of Justice, Vigil, Voyager; and receives research support for his laboratory from research grants from the National Institutes of Health, Cure Alzheimer’s Fund, Tau Consortium, and the JPB Foundation, and through sponsored research agreements from Abbvie, BMS, and Biogen. G.P.J. was the 2022 Past President of the American Society of Human Genetics. J.H.K. has been a consultant for Biogen. E.B.L. receives royalties from contributions to UpToDate. J.B.L. is a member of the Scientific Advisory Board of Vaxxinity and has received grant support from Biogen and GE Healthcare. A.I.L. is a founder of EmTheraPro. R.B.L. has received research support from the National Institutes of Health, the FDA, and the National Headache Foundation; serves as consultant, advisory board member, or has received honoraria or research support from AbbVie/Allergan, Amgen, Biohaven, Dr. Reddy’s Laboratories (Promius), electroCore, Eli Lilly, GlaxoSmithKline, Lundbeck, Merck, Novartis, Teva, Vector, and Vedanta Research; receives royalties from Wolff’s Headache, 8 th edition (Oxford University Press, 2009), and Informa; and holds stock in Biohaven and Manistee. D.C.M. receives NIH funding; is the inventor of the FCI-SF and the UAB Research Foundation (UABRF); owns the FCI-SF through copyright and trademark (FCAP); has previously received royalty and consulting income from UABRF licensed use and sale of the FCI-SF; and is currently a consultant on an unaffiliated NIH grant using the FCI-SF. A.V.M. is a council member of the Alzheimer's Association International Research Grants Program, on the steering committee of the Alzheimer's Disease Cooperative Study, and on the editorial boards of Alzheimer's & Dementia: Translational Research and Clinical Interventions and the Journal of Neuro-ophthalmology. S.I.M. is an employee of Pfizer, Inc.; the company did not influence the study design, analyses, or interpretation of the results presented in this manuscript. B.L.M. has received grant support from NIH, the Bluefield Project, and the Rainwater Charitable Foundation; has received royalties from books published by Cambridge University Press, Elsevier, Inc., Guilford Publications, Inc., Johns Hopkins Press, Oxford University Press and Taylor & Francis Group; has received honorarium for serving as a member of the Scientific Advisory Board of the Alzheimer’s Disease Research Center (ADRC) at Massachusetts General Hospital, Stanford University, and the University of Washington; and has received consulting fees from Genworth. J.W.M. receives compensation as Associate Editor for the journal Nutrition Reviews; and has received within the last 3 years consulting compensation from Church and Dwight, Inc.; and is a producer and seller of consumer goods including vitamin supplements. J.C.M is a consultant for clinical trials of antidementia drugs from Eli Lilly and Company, Biogen, and Janssen; is a consultant for the Barcelona Brain Research Center (BBRC) and the TS Srinivasan Advisory Board; is an advisory board member for the Cure Alzheimer’s Fund Research Strategy Council. S.E.O. has multiple pending and issued patents on blood biomarkers for detecting and precision medicine therapeutics in neurodegenerative diseases; and is a founding scientist of Cx Precision Medicine, Inc. and owns stock options. R.C.P. is chair for the data monitoring committee for Pfizer and Janssen Alzheimer Immunotherapy and is a consultant for GE Healthcare and Roche. W.W.P. is a co-inventor of and holds a patent for WO/2018/160496, related to the differentiation of human pluripotent stem cells into microglia. E.M.R. is a scientific advisor to Alzheon, Aural Analytics, Denali, Retromer Therapeutics, and Vaxxinity and a co-founder and advisor to ALZPath. J.M.R. receives research support from Avid Pharmaceuticals. R.N.R. is the Editor of JAMA Neurology. M.S. has received grants from the Icahn School of Medicine at Mount Sinai and the US Department of Veterans Affairs Veterans Health Administration. A.J.S. has received support from Avid Radiopharmaceuticals, a subsidiary of Eli Lilly (in-kind contribution of PET tracer precursor), is a member of scientific advisor boards for Bayer Oncology and Eisai and of the dementria advisory board and Siemens Medical Solutions USA, is a member of the National Heart, Lung, and Blood Institute MESA observational study monitoring board, and is part of the editorial office support as editor-in-chief for Brain Imaging and Behavior for Springer-Nature Publishing. J.A.S. has received consulting fees from AVID, Alnylam Pharmaceuticals, and Cerveau Technologies. L.S.S. has received personal fees from AC Immune, Athira, BioVie, Eli Lilly, Lundbeck, Merck, Neurim Ltd., Novo-Nordisk, Otsuka, Roche/Genentech within the past year; and research grants from Biogen, Eisai, and Eli Lilly and Company. W.W.S. serves as a paid consultant to Biogen Idec and has received grant support from NIH, the Association for Frontotemporal Degeneration, the Bluefield Project, the Rainwater Charitable Foundation, and the Chan-Zuckerberg Initiative. S.A.S. has received an unrestricted research grant from Mars, Inc. R.A.S. has received grants from the National Institutes of Health and the from Concussion Legacy Foundation; and received compensation from Biogen and Lundbeck; and has received royalties received from Psychological Assessment Resources for published neuropsychological tests; and has stock options as a member of the board of King Devick Technologies. D.L.S. has received research support from NIH and Eisai, has participated as a paid member of a DSMB or adjudication committee with Acadia, Avanir, Janssen, and Otsuka, and has received consulting fees from Avanir and NovoNordisk. R.E.T. has received patents for gamma-secretase modulators for exploring treatment of Alzheimer's disease. H.W. has received support from the TEVA speaker's bureau. T.S.W. is as a cofounder of revXon. C.B.W. has received royalties from UpTo Date for 2 chapters; has done legal consulting for the law firms of Abali, Milne, and Faegre Baker Daniels; is a consultant for Merck and Co; and does stroke adjudication for a National Institutes of Health clinical trial. L.A.F. has received institutional support from Mass Mutual Insurance. G.T., W.-P.L., Y.Y.L., P.P.K., J.B.M., J.A.P., R.M.S., X.Z., M.S.A., R.L.A., L.G.A., S.E.A., S.Asthana, C.T.B., R.C.B., L.L.B., S.B., T.G.B., J.T.B., D.Beekly, B.B., D.Bennett, T.D.B., D.A.B., B.F.B., J.D.B., A.L.B., J.M.B., J.D.Buxbaum, C.Cao, C.S.C., C.M.C., M.M.C., H.C.C., S.Craft, P.K.C., E.A.C., C.Cruchaga, M.L.C., P.L.D., C.D., J.C.D., M.Dick, D.W.D., R.S.D., R.D., N.E.-T., D.A.E., D.W.F., V.F., T.M.F., M.P.F., D.R.G., A.G., M.G., D.H.G., B.G., A.M.G., N.R.G.-R., R.C.G., H.H., O.Harari, J.Hardy, T.H., L.S.H., R.M.H., M.J.H., B.T.H., G.P.J., M.I.K., A.K., J.S.K., J.A.K., C.D.K., A.Khaleeq, N.W.K., J.H.K., W.K., E.B.L., J.B.L., A.I.L., A.P.L., R.B.L., M.W.L., O.L.L., K.L.L., C.G.L., M.E.F., D.C.M., E.R.M., D.C.Mash, E.M., A.V.M., W.C.M., A.C.M., M.Mesulam, B.L.M., C.A.M., J.W.M., T.J.M., J.C.M., S.Mukherjee, A.J.M., S.E.O., H.L.P., V.P., E.P., R.C.P., W.W.P., E.M.R., J.M.R., E.D.R., M.Rodriguear, R.N.R., M.S., A.J.S., J.A.S., L.S.S., W.W.S., S.A.S., R.A.S., S.M.S., D.S., R.E.T., D.W.T., V.M.VD., L.J.VE., J.M.V., B.N.V., E.M.W., T.S.W., C.B.W., S.G.Y., B.W.K., W.B., L.-S.W., L.A.F., J.L.H., R.M., M.A.P.-V., G.D.S., G.R.J., C.R., and A.C.N. have received grant funding from the National Institutes of Health, including from the National Institute on Aging and others. E.R. has received grant funding from the Canadian Institutes for Health Research.

Figures

**Fig. 1**
Genome-wide association findings across GSGs. Manhattan plot of cross-GSG meta-analyses of genome-wide associations estimated with covariate adjustment for age-at-onset (cases)/age-at-last-exam (controls), sex, and PCs for population substructure

**Fig. 2**
Cross-GSG comparison of multi-GSG GWAS associations. Shown are results for 18 loci associated with AD risk within and across multiple ancestries. Circle size corresponds to the significance level, − log₁₀(P-value), and circle color represents the significance threshold: red, P < 5 × 10⁻⁸; orange, 0.05 > P > 5 × 10⁻⁸; and blue, P > 0.05. Arrows represent the direction of effect of the risk allele, with “up” indicating increased risk and down decreased risk. Asterisks (“*”) indicate the association was found in *APOE* ε4-adjusted models

**Fig. 3**
Comparison of *LRRC4C* associations across GSGs. a Regional association plots and b a Forest plot for the novel *LRRC4C* locus cross-GSG and within-GSG associations with AD depicting all associations within ± 500 kb of the intergenic lead variant, rs12576934

**Fig. 4**
Comparison of *LHX5-AS1* associations across GSGs. a Regional association plots and b a Forest plot for the novel *LHX5-AS1* locus cross-GSG and within-GSG associations with AD depicting all associations within ± 500 kb of the intergenic lead variant, rs111486601

**Fig. 5**
Genome-wide association findings across GSGs with *APOE* ε4 adjustment. Manhattan plot of cross-GSG meta-analyses of genome-wide associations estimated with covariate adjustment for age-at-onset (cases)/age-at-last-exam (controls), sex, dosage of the *APOE* ε4 allele, and PCs for population substructure

See this image and copyright information in PMC

References

1. Alzheimer's Association. 2023 Alzheimer's disease facts and figures. Alzheimers Dement. 2023;19:1598–695. - PubMed
1. Kunkle BW, Grenier-Boley B, Sims R, Bis JC, Damotte V, Naj AC, Boland A, Vronskaya M, van der Lee SJ, Amlie-Wolf A, et al. Genetic meta-analysis of diagnosed Alzheimer’s disease identifies new risk loci and implicates Abeta, tau, immunity and lipid processing. Nat Genet. 2019;51:414–30. - PMC - PubMed
1. Jansen IE, Savage JE, Watanabe K, Bryois J, Williams DM, Steinberg S, Sealock J, Karlsson IK, Hagg S, Athanasiu L, et al. Genome-wide meta-analysis identifies new loci and functional pathways influencing Alzheimer’s disease risk. Nat Genet. 2019;51:404–13. - PMC - PubMed
1. Bellenguez C, Kucukali F, Jansen IE, Kleineidam L, Moreno-Grau S, Amin N, Naj AC, Campos-Martin R, Grenier-Boley B, Andrade V, et al. New insights into the genetic etiology of Alzheimer’s disease and related dementias. Nat Genet. 2022;54:412–36. - PMC - PubMed
1. Wightman DP, Jansen IE, Savage JE, Shadrin AA, Bahrami S, Holland D, Rongve A, Borte S, Winsvold BS, Drange OK, et al. A genome-wide association study with 1,126,563 individuals identifies new risk loci for Alzheimer’s disease. Nat Genet. 2021;53:1276–82. - PMC - PubMed

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Multi-ancestry genome-wide meta-analysis of 56,241 individuals identifies known and novel cross-population and ancestry-specific associations as novel risk loci for Alzheimer's disease

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Multi-ancestry genome-wide meta-analysis of 56,241 individuals identifies known and novel cross-population and ancestry-specific associations as novel risk loci for Alzheimer's disease

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