Updates are available Correction dated 2019-08-15
Click to view Correction: https://doi.org/10.1038/s41588-019-0495-7
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Authors
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Kunkle, Brian W. https://orcid.org/0000-0002-9515-5157
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Grenier-Boley, Benjamin
Sims, Rebecca
Bis, Joshua C.
Damotte, Vincent
Naj, Adam C.
Boland, Anne
Vronskaya, Maria
van der Lee, Sven J. https://orcid.org/0000-0003-1606-8643
Amlie-Wolf, Alexandre https://orcid.org/0000-0002-2073-1519
Bellenguez, Céline https://orcid.org/0000-0002-1240-7874
Frizatti, Aura
Chouraki, Vincent
Martin, Eden R.
Sleegers, Kristel https://orcid.org/0000-0002-0283-2332
Badarinarayan, Nandini
Jakobsdottir, Johanna
Hamilton-Nelson, Kara L.
Moreno-Grau, Sonia
Olaso, Robert
Raybould, Rachel
Chen, Yuning https://orcid.org/0000-0002-7358-7055
Kuzma, Amanda B.
Hiltunen, Mikko
Morgan, Taniesha
Ahmad, Shahzad
Vardarajan, Badri N.
Epelbaum, Jacques
Hoffmann, Per
Boada, Merce
Beecham, Gary W.
Garnier, Jean-Guillaume
Harold, Denise
Fitzpatrick, Annette L.
Valladares, Otto
Moutet, Marie-Laure
Gerrish, Amy
Smith, Albert V.
Qu, Liming
Bacq, Delphine
Denning, Nicola
Jian, Xueqiu
Zhao, Yi
Del Zompo, Maria
Fox, Nick C.
Choi, Seung-Hoan
Mateo, Ignacio
Hughes, Joseph T.
Adams, Hieab H. https://orcid.org/0000-0003-3687-2508
Malamon, John
Sanchez-Garcia, Florentino
Patel, Yogen
Brody, Jennifer A. https://orcid.org/0000-0001-8509-148X
Dombroski, Beth A.
Naranjo, Maria Candida Deniz
Daniilidou, Makrina
Eiriksdottir, Gudny
Mukherjee, Shubhabrata
Wallon, David
Uphill, James
Aspelund, Thor https://orcid.org/0000-0002-7998-5433
Cantwell, Laura B.
Garzia, Fabienne
Galimberti, Daniela
Hofer, Edith
Butkiewicz, Mariusz
Fin, Bertrand
Scarpini, Elio
Sarnowski, Chloe
Bush, Will S. https://orcid.org/0000-0002-9729-6519
Meslage, Stéphane
Kornhuber, Johannes
White, Charles C.
Song, Yuenjoo
Barber, Robert C. https://orcid.org/0000-0001-6857-0286
Engelborghs, Sebastiaan
Sordon, Sabrina
Voijnovic, Dina
Adams, Perrie M.
Vandenberghe, Rik
Mayhaus, Manuel
Cupples, L. Adrienne https://orcid.org/0000-0003-0273-7965
Albert, Marilyn S.
De Deyn, Peter P. https://orcid.org/0000-0002-2228-2964
Gu, Wei
Himali, Jayanadra J. https://orcid.org/0000-0003-1391-9481
Beekly, Duane
Squassina, Alessio https://orcid.org/0000-0001-7415-7607
Hartmann, Annette M.
Orellana, Adelina
Blacker, Deborah
Rodriguez-Rodriguez, Eloy
Lovestone, Simon
Garcia, Melissa E.
Doody, Rachelle S.
Munoz-Fernadez, Carmen
Sussams, Rebecca
Lin, Honghuang https://orcid.org/0000-0003-3043-3942
Fairchild, Thomas J.
Benito, Yolanda A.
Holmes, Clive
Karamujić-Čomić, Hata
Frosch, Matthew P.
Thonberg, Hakan
Maier, Wolfgang
Roshchupkin, Gennady https://orcid.org/0000-0002-3403-2313
Ghetti, Bernardino https://orcid.org/0000-0002-1842-8019
Giedraitis, Vilmantas https://orcid.org/0000-0003-3423-2021
Kawalia, Amit
Li, Shuo https://orcid.org/0000-0003-2331-2448
Huebinger, Ryan M.
Kilander, Lena
Moebus, Susanne
Hernández, Isabel
Kamboh, M. Ilyas
Brundin, RoseMarie
Turton, James
Yang, Qiong https://orcid.org/0000-0002-3658-1375
Katz, Mindy J.
Concari, Letizia
Lord, Jenny
Beiser, Alexa S.
Keene, C. Dirk
Helisalmi, Seppo
Kloszewska, Iwona
Kukull, Walter A.
Koivisto, Anne Maria
Lynch, Aoibhinn
Tarraga, Lluís
Larson, Eric B.
Haapasalo, Annakaisa
Lawlor, Brian
Mosley, Thomas H.
Lipton, Richard B.
Solfrizzi, Vincenzo
Gill, Michael https://orcid.org/0000-0003-0206-5337
Longstreth, W. T. Jr
Montine, Thomas J.
Frisardi, Vincenza
Diez-Fairen, Monica
Rivadeneira, Fernando https://orcid.org/0000-0001-9435-9441
Petersen, Ronald C.
Deramecourt, Vincent
Alvarez, Ignacio
Salani, Francesca
Ciaramella, Antonio
Boerwinkle, Eric
Reiman, Eric M.
Fievet, Nathalie
Rotter, Jerome I.
Reisch, Joan S.
Hanon, Olivier
Cupidi, Chiara
Andre Uitterlinden, A. G.
Royall, Donald R.
Dufouil, Carole
Maletta, Raffaele Giovanni
de Rojas, Itziar
Sano, Mary
Brice, Alexis
Cecchetti, Roberta
George-Hyslop, Peter St
Ritchie, Karen
Tsolaki, Magda https://orcid.org/0000-0002-2072-8010
Tsuang, Debby W.
Dubois, Bruno
Craig, David
Wu, Chuang-Kuo
Soininen, Hilkka
Avramidou, Despoina
Albin, Roger L.
Fratiglioni, Laura
Germanou, Antonia
Apostolova, Liana G.
Keller, Lina
Koutroumani, Maria
Arnold, Steven E.
Panza, Francesco
Gkatzima, Olymbia
Asthana, Sanjay
Hannequin, Didier
Whitehead, Patrice
Atwood, Craig S. https://orcid.org/0000-0001-5040-1679
Caffarra, Paolo
Hampel, Harald
Quintela, Inés
Carracedo, Ángel
Lannfelt, Lars
Rubinsztein, David C. https://orcid.org/0000-0001-5002-5263
Barnes, Lisa L.
Pasquier, Florence https://orcid.org/0000-0001-9880-9788
Frölich, Lutz
Barral, Sandra
McGuinness, Bernadette
Beach, Thomas G.
Johnston, Janet A.
Becker, James T. https://orcid.org/0000-0003-4425-4726
Passmore, Peter
Bigio, Eileen H.
Schott, Jonathan M. https://orcid.org/0000-0003-2059-024X
Bird, Thomas D.
Warren, Jason D.
Boeve, Bradley F.
Lupton, Michelle K. https://orcid.org/0000-0002-7274-7299
Bowen, James D. https://orcid.org/0000-0002-4258-250X
Proitsi, Petra
Boxer, Adam
Powell, John F.
Burke, James R. https://orcid.org/0000-0002-3408-7787
Kauwe, John S. K.
Burns, Jeffrey M.
Mancuso, Michelangelo https://orcid.org/0000-0003-2738-8562
Buxbaum, Joseph D.
Bonuccelli, Ubaldo
Cairns, Nigel J.
McQuillin, Andrew https://orcid.org/0000-0003-1567-2240
Cao, Chuanhai
Livingston, Gill
Carlson, Chris S.
Bass, Nicholas J.
Carlsson, Cynthia M.
Hardy, John
Carney, Regina M.
Bras, Jose
Carrasquillo, Minerva M.
Guerreiro, Rita
Allen, Mariet
Chui, Helena C.
Fisher, Elizabeth https://orcid.org/0000-0003-2850-9936
Masullo, Carlo
Crocco, Elizabeth A.
DeCarli, Charles
Bisceglio, Gina
Dick, Malcolm
Ma, Li
Duara, Ranjan
Graff-Radford, Neill R.
Evans, Denis A.
Hodges, Angela
Faber, Kelley M.
Scherer, Martin
Fallon, Kenneth B.
Riemenschneider, Matthias
Fardo, David W. https://orcid.org/0000-0002-7207-4696
Heun, Reinhard
Farlow, Martin R.
Kölsch, Heike
Ferris, Steven
Leber, Markus
Foroud, Tatiana M.
Heuser, Isabella
Galasko, Douglas R.
Giegling, Ina
Gearing, Marla
Hüll, Michael
Geschwind, Daniel H.
Gilbert, John R.
Morris, John
Green, Robert C.
Mayo, Kevin
Growdon, John H.
Feulner, Thomas
Hamilton, Ronald L. https://orcid.org/0000-0002-6802-1102
Harrell, Lindy E.
Drichel, Dmitriy
Honig, Lawrence S.
Cushion, Thomas D.
Huentelman, Matthew J.
Hollingworth, Paul
Hulette, Christine M.
Hyman, Bradley T.
Marshall, Rachel
Jarvik, Gail P.
Meggy, Alun
Abner, Erin
Menzies, Georgina E. https://orcid.org/0000-0002-6600-6507
Jin, Lee-Way
Leonenko, Ganna
Real, Luis M.
Jun, Gyungah R.
Baldwin, Clinton T.
Grozeva, Detelina
Karydas, Anna
Russo, Giancarlo
Kaye, Jeffrey A.
Kim, Ronald
Jessen, Frank
Kowall, Neil W.
Vellas, Bruno
Kramer, Joel H.
Vardy, Emma
LaFerla, Frank M.
Jöckel, Karl-Heinz
Lah, James J.
Dichgans, Martin
Leverenz, James B.
Mann, David
Levey, Allan I.
Pickering-Brown, Stuart https://orcid.org/0000-0003-1561-6054
Lieberman, Andrew P.
Klopp, Norman
Lunetta, Kathryn L. https://orcid.org/0000-0002-9268-810X
Wichmann, H-Erich
Lyketsos, Constantine G.
Morgan, Kevin https://orcid.org/0000-0002-8217-2396
Marson, Daniel C.
Brown, Kristelle
Martiniuk, Frank
Medway, Christopher
Mash, Deborah C.
Nöthen, Markus M.
Masliah, Eliezer
Hooper, Nigel M. https://orcid.org/0000-0002-5811-3484
McCormick, Wayne C.
Daniele, Antonio
McCurry, Susan M.
Bayer, Anthony
McDavid, Andrew N. https://orcid.org/0000-0002-6581-1213
Gallacher, John https://orcid.org/0000-0002-2394-5299
McKee, Ann C.
van den Bussche, Hendrik
Mesulam, Marsel
Brayne, Carol
Miller, Bruce L.
Riedel-Heller, Steffi
Miller, Carol A.
Miller, Joshua W.
Al-Chalabi, Ammar https://orcid.org/0000-0002-4924-7712
Morris, John C.
Shaw, Christopher E.
Myers, Amanda J.
Wiltfang, Jens
O’Bryant, Sid
Olichney, John M.
Alvarez, Victoria
Parisi, Joseph E.
Singleton, Andrew B.
Paulson, Henry L.
Collinge, John
Perry, William R.
Mead, Simon
Peskind, Elaine
Cribbs, David H.
Rossor, Martin
Pierce, Aimee
Ryan, Natalie S.
Poon, Wayne W. https://orcid.org/0000-0003-0625-3968
Nacmias, Benedetta
Potter, Huntington
Sorbi, Sandro
Quinn, Joseph F.
Sacchinelli, Eleonora https://orcid.org/0000-0002-7648-5126
Raj, Ashok
Spalletta, Gianfranco
Raskind, Murray
Caltagirone, Carlo
Bossù, Paola
Orfei, Maria Donata
Reisberg, Barry
Clarke, Robert https://orcid.org/0000-0002-9802-8241
Reitz, Christiane
Smith, A David
Ringman, John M.
Warden, Donald
Roberson, Erik D. https://orcid.org/0000-0002-1810-9763
Wilcock, Gordon
Rogaeva, Ekaterina
Bruni, Amalia Cecilia
Rosen, Howard J.
Gallo, Maura
Rosenberg, Roger N.
Ben-Shlomo, Yoav
Sager, Mark A.
Mecocci, Patrizia
Saykin, Andrew J. https://orcid.org/0000-0002-1376-8532
Pastor, Pau https://orcid.org/0000-0002-7493-8777
Cuccaro, Michael L.
Vance, Jeffery M.
Schneider, Julie A.
Schneider, Lori S.
Slifer, Susan
Seeley, William W.
Smith, Amanda G.
Sonnen, Joshua A.
Spina, Salvatore
Stern, Robert A.
Swerdlow, Russell H.
Tang, Mitchell
Tanzi, Rudolph E. https://orcid.org/0000-0002-7032-1454
Trojanowski, John Q.
Troncoso, Juan C.
Van Deerlin, Vivianna M.
Van Eldik, Linda J.
Vinters, Harry V.
Vonsattel, Jean Paul
Weintraub, Sandra
Welsh-Bohmer, Kathleen A.
Wilhelmsen, Kirk C.
Williamson, Jennifer
Wingo, Thomas S. https://orcid.org/0000-0002-7679-6282
Woltjer, Randall L.
Wright, Clinton B.
Yu, Chang-En
Yu, Lei
Saba, Yasaman
Pilotto, Alberto
Bullido, Maria J. https://orcid.org/0000-0002-6477-1117
Peters, Oliver
Crane, Paul K.
Bennett, David
Bosco, Paola
Coto, Eliecer
Boccardi, Virginia
De Jager, Phil L. https://orcid.org/0000-0002-8057-2505
Lleo, Alberto
Warner, Nick
Lopez, Oscar L.
Ingelsson, Martin
Deloukas, Panagiotis https://orcid.org/0000-0001-9251-070X
Cruchaga, Carlos https://orcid.org/0000-0002-0276-2899
Graff, Caroline
Gwilliam, Rhian
Fornage, Myriam
Goate, Alison M. https://orcid.org/0000-0002-0576-2472
Sanchez-Juan, Pascual
Kehoe, Patrick G.
Amin, Najaf
Ertekin-Taner, Nilifur
Berr, Claudine https://orcid.org/0000-0001-5254-7655
Debette, Stéphanie
Love, Seth
Launer, Lenore J.
Younkin, Steven G.
Dartigues, Jean-Francois
Corcoran, Chris
Ikram, M. Arfan https://orcid.org/0000-0003-0372-8585
Dickson, Dennis W. https://orcid.org/0000-0001-7189-7917
Nicolas, Gael
Campion, Dominique
Tschanz, JoAnn
Schmidt, Helena
Hakonarson, Hakon
Clarimon, Jordi
Munger, Ron
Schmidt, Reinhold
Farrer, Lindsay A.
Van Broeckhoven, Christine https://orcid.org/0000-0003-0183-7665
C. O’Donovan, Michael https://orcid.org/0000-0001-7073-2379
DeStefano, Anita L.
Jones, Lesley https://orcid.org/0000-0002-3007-4612
Haines, Jonathan L.
Deleuze, Jean-Francois
Owen, Michael J. https://orcid.org/0000-0003-4798-0862
Gudnason, Vilmundur
Mayeux, Richard
Escott-Price, Valentina https://orcid.org/0000-0003-1784-5483
Psaty, Bruce M.
Ramirez, Alfredo https://orcid.org/0000-0003-4991-763X
Wang, Li-San
Ruiz, Agustin
van Duijn, Cornelia M.
Holmans, Peter A.
Seshadri, Sudha
Williams, Julie
Amouyel, Phillippe https://orcid.org/0000-0001-9088-234X
Schellenberg, Gerard D.
Lambert, Jean-Charles https://orcid.org/0000-0003-0829-7817
Pericak-Vance, Margaret A. https://orcid.org/0000-0001-7283-8804
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Article History
Received: 12 March 2018
Accepted: 22 January 2019
First Online: 28 February 2019
Change Date: 15 August 2019
Change Type: Correction
Change Details: An amendment to this paper has been published and can be accessed via a link at the top of the paper.
Competing interests
: D. Blacker is a consultant for Biogen, Inc. R.C.P. is a consultant for Roche, Inc.; Merck, Inc.; Genentech, Inc.; Biogen, Inc.; GE Healthcare; and Eisai, Inc. A.R.W. is a former employee and stockholder of Pfizer, Inc., and a current employee of the Perelman School of Medicine at the University of Pennsylvania Orphan Disease Center in partnership with the Loulou. A.M.G. is a member of the scientific advisory board for Denali Therapeutics. N.E.-T. is a consultant for Cytox. J.Hardy holds a collaborative grant with Cytox cofunded by the Department of Business (Biz). F.J. acts as a consultant for Novartis, Eli Lilly, Nutricia, MSD, Roche and Piramal. Neither J.M. nor his family owns stock or has equity interest (outside of mutual funds or other externally directed accounts) in any pharmaceutical or biotechnology company. J.M. is currently participating in clinical trials of antidementia drugs from Eli Lilly and Company, Biogen and Janssen. J.M. serves as a consultant for Lilly USA. He receives research support from Eli Lilly/Avid Radiopharmaceuticals and is funded by NIH grant nos. P50AG005681, P01AG003991, P01AG026276 and UF01AG032438. C.Cruchaga receives research support from Biogen, EISAI, Alector and Parabon. The funders of the study had no role in the collection, analysis or interpretation of data; in the writing of the report; or in the decision to submit the paper for publication. C.Cruchaga is a member of the advisory board of ADx Healthcare. M.R.F. receives grant/research support from AbbVie, Accera, ADCS Posiphen, Biogen, Eisai, Eli Lilly, Genentech, Novartis and Suven Life Sciences, Ltd. He is a consultant/advisory board/DSMB board member for Accera, Avanir, AZTherapies, Cognition Therapeutics, Cortexyme, Eli Lilly & Company, Longeveron, Medavante, Merck and Co. Inc., Otsuka Pharmaceutical, Proclara (formerly Neurophage Pharmaceuticals), Neurotrope Biosciences, Takeda, vTv Therapeutics and Zhejian Hisun Pharmaceuticals. He has a transgenic mouse model patent that is licensed to Elan. R.A.S. receives consulting fees as a member of the Alzheimer’s Disease Advisory Board, Biogen; and as member of the Executive Committee for AZD3293 Alzheimer’s Disease Studies, Eli Lilly. R.B.L. receives consulting fees from Merch, Inc. E.M.R. receives grant funding from several NIH grant and research contracts with Genentech/Roche, Novartis/Amgen and Avid/Lilly. He is a compensated scientific advisor to Alkahest, Alzheon, Aural Analytics, Denali, Takeda and Zinfandel. He is an advisor to Roche and Roche Diagnostics, which reimburse his expenses only. T.G.B. has research support/contracts from the National Institutes of Health, State of Arizona, Michael J Fox Foundation, Avid Radiopharmaceuticals, Navida Biopharmaceuticals and Aprinoia Therapeutics. He is an advisory board member with Vivid Genomics and has consultancy work with Roche Diagnostics. A.G.S. conducts multiple industry-funded clinical trials, but all funds go to her academic institution. They have current (within last 12 months) research contracts with Eli Lilly, Novartis, Roche, Janssen, AbbVie, Biogen, NeuroEM, Suven and Merck. She does not receive personal compensations from these organizations. G.D.S. is a consultant for Biogen, Inc. J.M.B. is participating in clinical trials of antidementia drugs for Eli Lilly, Toyama Chemical Company, Merck, Biogen, AbbVie, vTv Therapeutics, Janssen and Roche. He has received research grants from Eli Lilly, Avid Radiopharmaceuticals and Astra Zeneca. He is a consultant for Stage 2 Innovations. L.F. is a consultant for Allergan, Eli Lilly, Avraham Pharmaceuticals, Axon Neuroscience, Axovant, Biogen, Boehringer Ingelheim, Eisai, Functional Neuromodulation, Lundbeck, MerckSharpe & Dohme, Novartis, Pfizer, Pharnext, Roche and Schwabe Pharma. M.B has consulted as an advisory board member for Araclon, Grifols, Lilly, Nutricia, Roche and Servier. She received fees for lectures and funds for research from Araclon, Grifols, Nutricia, Roche and Servier. She has not received personal compensations from these organizations. A.Ruiz has consulted for Grifols and Landsteiner Genmed. He received fees for lectures or funds for research and/or reimbursement of expenses for congresses attendance from Araclon and Grifols. He has not received personal compensations from these organizations. O.P. acts as a consultant for Roche and Biogen, Inc. He is currently participating in clinical trials of antidementia drugs from Novartis, Genentech, Roche and Pharmatrophix. B.T.H. is a consultant for Aztherapy, Biogen, Calico, Ceregene, Genentech, Lilly, Neurophage, Novartis and Takeda, and receives research support from Abbvie, Amgen, Deanli, Fidelity Biosciences, General Electric, Lilly, Merck, Sangamo and Spark therapeutics. BTH owns Novartis stock. H.Hampel serves as Senior Associate Editor for the Journal Alzheimer’s & Dementia; he received lecture fees from Biogen and Roche, research grants from Pfizer, Avid and MSD AVENIR (paid to the institution), travel funding from Functional Neuromodulation, Axovant, Eli Lilly and company, Takeda and Zinfandel, GE Healthcare and Oryzon Genomics, consultancy fees from Jung Diagnostics, Cytox Ltd., Axovant, Anavex, Takeda and Zinfandel, GE Healthcare, Oryzon Genomics and Functional Neuromodulation, and participated in scientific advisory boards of Functional Neuromodulation, Axovant, Eli Lilly and company, Cytox Ltd., GE Healthcare, Takeda and Zinfandel, Oryzon Genomics and Roche Diagnostica. Harald Hampel is a co-inventor on numerous patents relating to biomarker measurement but has received no royalties from these patents. A.A.-C. has consultancies for GSK, Cytokinetics, Biogen Idec, Treeway Inc, Chronos Therapeutics, OrionPharma and Mitsubishi-Tanabe Pharma, and was Chief Investigator for commercial clinical trials run by OrionPharma and Cytokinetics.