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CORTICAL GENETICS

The genetic architecture of the human cerebral cortex

Katrina L. Grasby https://orcid.org/0000-0001-8539-0228 [email protected], Neda Jahanshad https://orcid.org/0000-0003-4401-8950 [email protected], Jodie N. Painter https://orcid.org/0000-0002-4992-2458, Lucía Colodro-Conde https://orcid.org/0000-0002-9004-364X, Janita Bralten https://orcid.org/0000-0003-1440-8675, Derrek P. Hibar, Penelope A. Lind https://orcid.org/0000-0002-3887-2598, Fabrizio Pizzagalli, Christopher R. K. Ching, Mary Agnes B. McMahon https://orcid.org/0000-0002-1516-2462, Natalia Shatokhina, Leo C. P. Zsembik https://orcid.org/0000-0003-2609-5637, Sophia I. Thomopoulos, Alyssa H. Zhu https://orcid.org/0000-0003-0083-5107, Lachlan T. Strike https://orcid.org/0000-0003-2885-5898, Ingrid Agartz https://orcid.org/0000-0002-9839-5391, Saud Alhusaini, Marcio A. A. Almeida https://orcid.org/0000-0002-8114-3892, Dag Alnæs, Inge K. Amlien https://orcid.org/0000-0002-8508-9088, Micael Andersson https://orcid.org/0000-0003-4743-6365, Tyler Ard https://orcid.org/0000-0002-5123-1226, Nicola J. Armstrong https://orcid.org/0000-0002-4477-293X, Allison Ashley-Koch, Joshua R. Atkins https://orcid.org/0000-0003-0821-1112, Manon Bernard https://orcid.org/0000-0002-7234-1782, Rachel M. Brouwer https://orcid.org/0000-0002-7466-1544, Elizabeth E. L. Buimer https://orcid.org/0000-0002-8143-3160, Robin Bülow, Christian Bürger, Dara M. Cannon, Mallar Chakravarty, Qiang Chen https://orcid.org/0000-0003-0373-4459, Joshua W. Cheung https://orcid.org/0000-0002-3976-5073, Baptiste Couvy-Duchesne https://orcid.org/0000-0002-0719-9302, Anders M. Dale https://orcid.org/0000-0002-6126-2966, Shareefa Dalvie, Tânia K. de Araujo https://orcid.org/0000-0003-3056-3894, Greig I. de Zubicaray https://orcid.org/0000-0003-4506-0579, Sonja M. C. de Zwarte, Anouk den Braber, Nhat Trung Doan, Katharina Dohm https://orcid.org/0000-0002-9305-9946, Stefan Ehrlich https://orcid.org/0000-0003-2132-4445, Hannah-Ruth Engelbrecht https://orcid.org/0000-0002-6627-4114, Susanne Erk, Chun Chieh Fan, Iryna O. Fedko https://orcid.org/0000-0003-1962-916X, Sonya F. Foley, Judith M. Ford https://orcid.org/0000-0002-6500-6548, Masaki Fukunaga https://orcid.org/0000-0003-1010-2644, Melanie E. Garrett https://orcid.org/0000-0003-2302-6150, Tian Ge, Sudheer Giddaluru, Aaron L. Goldman, Melissa J. Green https://orcid.org/0000-0002-9361-4874, Nynke A. Groenewold, Dominik Grotegerd https://orcid.org/0000-0001-8718-0128, Tiril P. Gurholt https://orcid.org/0000-0002-1272-7616, Boris A. Gutman https://orcid.org/0000-0001-5388-897X, Narelle K. Hansell https://orcid.org/0000-0002-8229-9741, Mathew A. Harris https://orcid.org/0000-0002-1135-4141, Marc B. Harrison https://orcid.org/0000-0002-2742-241X, Courtney C. Haswell https://orcid.org/0000-0003-0562-9855, Michael Hauser https://orcid.org/0000-0003-2667-5420, Stefan Herms https://orcid.org/0000-0002-2786-8200, Dirk J. Heslenfeld, New Fei Ho, David Hoehn https://orcid.org/0000-0002-2501-2468, Per Hoffmann https://orcid.org/0000-0002-6573-983X, Laurena Holleran https://orcid.org/0000-0002-8354-3670, Martine Hoogman https://orcid.org/0000-0002-1261-7628, Jouke-Jan Hottenga https://orcid.org/0000-0002-5668-2368, Masashi Ikeda https://orcid.org/0000-0001-6237-2449, Deborah Janowitz https://orcid.org/0000-0003-4324-9298, Iris E. Jansen https://orcid.org/0000-0003-1901-8131, Tianye Jia https://orcid.org/0000-0001-5399-2953, Christiane Jockwitz https://orcid.org/0000-0001-9332-7982, Ryota Kanai https://orcid.org/0000-0002-0186-2687, Sherif Karama https://orcid.org/0000-0002-9379-7180, Dalia Kasperaviciute, Tobias Kaufmann https://orcid.org/0000-0002-4003-1018, Sinead Kelly, Masataka Kikuchi https://orcid.org/0000-0002-9562-841X, Marieke Klein https://orcid.org/0000-0001-8784-5679, Michael Knapp, Annchen R. Knodt https://orcid.org/0000-0001-6501-8947, Bernd Krämer https://orcid.org/0000-0002-1145-9103, Max Lam, Thomas M. Lancaster, Phil H. Lee https://orcid.org/0000-0003-1770-3100, Tristram A. Lett, Lindsay B. Lewis https://orcid.org/0000-0002-3056-4152, Iscia Lopes-Cendes https://orcid.org/0000-0002-6221-6822, Michelle Luciano https://orcid.org/0000-0002-7306-3008, Fabio Macciardi https://orcid.org/0000-0003-0537-4266, Andre F. Marquand https://orcid.org/0000-0001-5903-203X, Samuel R. Mathias, Tracy R. Melzer https://orcid.org/0000-0003-0621-212X, Yuri Milaneschi, Nazanin Mirza-Schreiber https://orcid.org/0000-0003-0836-8267, Jose C. V. Moreira https://orcid.org/0000-0002-9527-7349, Thomas W. Mühleisen https://orcid.org/0000-0002-6057-5952, Bertram Müller-Myhsok, Pablo Najt https://orcid.org/0000-0002-4263-3880, Soichiro Nakahara, Kwangsik Nho, Loes M. Olde Loohuis https://orcid.org/0000-0002-3327-7837, Dimitri Papadopoulos Orfanos https://orcid.org/0000-0002-1242-8990, John F. Pearson https://orcid.org/0000-0001-5607-4517, Toni L. Pitcher https://orcid.org/0000-0003-4661-5038, Benno Pütz, Yann Quidé https://orcid.org/0000-0002-8569-7139, Anjanibhargavi Ragothaman https://orcid.org/0000-0001-9243-9368, Faisal M. Rashid, William R. Reay https://orcid.org/0000-0001-7689-2453, Ronny Redlich, Céline S. Reinbold https://orcid.org/0000-0002-3394-3158, Jonathan Repple https://orcid.org/0000-0003-1379-9491, Geneviève Richard https://orcid.org/0000-0001-6475-2576, Brandalyn C. Riedel https://orcid.org/0000-0001-9047-7512, Shannon L. Risacher, Cristiane S. Rocha https://orcid.org/0000-0001-5678-2070, Nina Roth Mota https://orcid.org/0000-0003-3504-759X, Lauren Salminen https://orcid.org/0000-0002-0684-9991, Arvin Saremi https://orcid.org/0000-0003-2395-8249, Andrew J. Saykin https://orcid.org/0000-0002-1376-8532, Fenja Schlag https://orcid.org/0000-0002-4867-2530, Lianne Schmaal, Peter R. Schofield https://orcid.org/0000-0003-2967-9662, Rodrigo Secolin https://orcid.org/0000-0002-2485-9560, Chin Yang Shapland https://orcid.org/0000-0002-5797-1241, Li Shen https://orcid.org/0000-0002-5443-0503, Jean Shin https://orcid.org/0000-0001-7353-5178, Elena Shumskaya https://orcid.org/0000-0003-4383-3988, Ida E. Sønderby, Emma Sprooten https://orcid.org/0000-0002-1691-9105, Katherine E. Tansey, Alexander Teumer, Anbupalam Thalamuthu https://orcid.org/0000-0002-7114-1260, Diana Tordesillas-Gutiérrez, Jessica A. Turner https://orcid.org/0000-0003-0076-8434, Anne Uhlmann, Costanza Ludovica Vallerga https://orcid.org/0000-0001-9493-6800, Dennis van der Meer, Marjolein M. J. van Donkelaar, Liza van Eijk https://orcid.org/0000-0001-8254-1019, Theo G. M. van Erp https://orcid.org/0000-0002-2465-2797, Neeltje E. M. van Haren https://orcid.org/0000-0002-3090-7653, Daan van Rooij https://orcid.org/0000-0002-6681-4344, Marie-José van Tol https://orcid.org/0000-0002-9260-5490, Jan H. Veldink https://orcid.org/0000-0001-5572-9657, Ellen Verhoef https://orcid.org/0000-0002-8315-1095, Esther Walton https://orcid.org/0000-0002-0935-2200, Mingyuan Wang https://orcid.org/0000-0003-1946-1942, Yunpeng Wang https://orcid.org/0000-0001-9831-1090, Joanna M. Wardlaw https://orcid.org/0000-0002-9812-6642, Wei Wen https://orcid.org/0000-0003-2753-3870, Lars T. Westlye https://orcid.org/0000-0001-8644-956X, Christopher D. Whelan, Stephanie H. Witt https://orcid.org/0000-0002-1571-1468, Katharina Wittfeld https://orcid.org/0000-0003-4383-5043, Christiane Wolf https://orcid.org/0000-0002-6587-7407, Thomas Wolfers https://orcid.org/0000-0002-7693-0621, Jing Qin Wu https://orcid.org/0000-0002-2052-3875, Clarissa L. Yasuda https://orcid.org/0000-0001-9084-7173, Dario Zaremba, Zuo Zhang https://orcid.org/0000-0002-6752-3434, Marcel P. Zwiers, Eric Artiges https://orcid.org/0000-0003-4461-7646, Amelia A. Assareh, Rosa Ayesa-Arriola https://orcid.org/0000-0003-0570-5352, Aysenil Belger https://orcid.org/0000-0003-2687-1966, Christine L. Brandt, Gregory G. Brown, Sven Cichon https://orcid.org/0000-0002-9475-086X, Joanne E. Curran https://orcid.org/0000-0002-6898-155X, Gareth E. Davies, Franziska Degenhardt, Michelle F. Dennis https://orcid.org/0000-0003-0734-7448, Bruno Dietsche, Srdjan Djurovic https://orcid.org/0000-0002-8140-8061, Colin P. Doherty https://orcid.org/0000-0002-8869-8567, Ryan Espiritu, Daniel Garijo https://orcid.org/0000-0003-0454-7145, Yolanda Gil, Penny A. Gowland https://orcid.org/0000-0002-4900-4817, Robert C. Green https://orcid.org/0000-0001-8472-0424, Alexander N. Häusler https://orcid.org/0000-0003-3506-7566, Walter Heindel, Beng-Choon Ho https://orcid.org/0000-0003-3976-1555, Wolfgang U. Hoffmann https://orcid.org/0000-0002-6359-8797, Florian Holsboer https://orcid.org/0000-0002-2640-9336, Georg Homuth https://orcid.org/0000-0001-6839-0605, Norbert Hosten https://orcid.org/0000-0002-4149-5666, Clifford R. JackJr. https://orcid.org/0000-0001-7916-622X, MiHyun Jang https://orcid.org/0000-0001-5638-2237, Andreas Jansen https://orcid.org/0000-0002-3140-8207, Nathan A. Kimbrel, Knut Kolskår, Sanne Koops, Axel Krug https://orcid.org/0000-0002-0564-2497, Kelvin O. Lim https://orcid.org/0000-0002-2390-7268, Jurjen J. Luykx https://orcid.org/0000-0002-6439-2774, Daniel H. Mathalon https://orcid.org/0000-0001-6090-4974, Karen A. Mather, Venkata S. Mattay https://orcid.org/0000-0002-3829-3369, Sarah Matthews, Jaqueline Mayoral Van Son, Sarah C. McEwen https://orcid.org/0000-0001-6425-6126, Ingrid Melle https://orcid.org/0000-0002-9783-548X, Derek W. Morris https://orcid.org/0000-0002-3413-570X, Bryon A. Mueller https://orcid.org/0000-0003-2429-0391, Matthias Nauck, Jan E. Nordvik https://orcid.org/0000-0003-0981-648X, Markus M. Nöthen https://orcid.org/0000-0002-8770-2464, Daniel S. O’Leary https://orcid.org/0000-0003-1959-6521, Nils Opel, Marie-Laure Paillère Martinot, G. Bruce Pike, Adrian Preda https://orcid.org/0000-0003-3373-2438, Erin B. Quinlan, Paul E. Rasser https://orcid.org/0000-0003-3482-721X, Varun Ratnakar https://orcid.org/0000-0001-9381-705X, Simone Reppermund, Vidar M. Steen https://orcid.org/0000-0001-6440-3966, Paul A. Tooney, Fábio R. Torres https://orcid.org/0000-0001-6550-6887, Dick J. Veltman https://orcid.org/0000-0001-8971-4250, James T. Voyvodic https://orcid.org/0000-0003-1072-6158, Robert Whelan https://orcid.org/0000-0002-2790-7281, Tonya White, Hidenaga Yamamori https://orcid.org/0000-0003-2100-1881, Hieab H. H. Adams https://orcid.org/0000-0003-3687-2508, Joshua C. Bis, Stephanie Debette, Charles Decarli, Myriam Fornage, Vilmundur Gudnason, Edith Hofer, M. Arfan Ikram, Lenore Launer, W. T. Longstreth, Oscar L. Lopez https://orcid.org/0000-0002-8546-8256, Bernard Mazoyer https://orcid.org/0000-0003-0970-2837, Thomas H. Mosley https://orcid.org/0000-0003-3343-1352, Gennady V. Roshchupkin, Claudia L. Satizabal https://orcid.org/0000-0002-1115-4430, Reinhold Schmidt, Sudha Seshadri https://orcid.org/0000-0001-6135-2622, Qiong Yang, Alzheimer’s Disease Neuroimaging Initiative, CHARGE Consortium, EPIGEN Consortium, IMAGEN Consortium, SYS Consortium, Parkinson’s Progression Markers Initiative, Marina K. M. Alvim https://orcid.org/0000-0002-6780-0761, David Ames, Tim J. Anderson https://orcid.org/0000-0002-1116-3839, Ole A. Andreassen https://orcid.org/0000-0002-4461-3568, Alejandro Arias-Vasquez, Mark E. Bastin https://orcid.org/0000-0002-0490-0845, Bernhard T. Baune, Jean C. Beckham https://orcid.org/0000-0001-8746-8949, John Blangero, Dorret I. Boomsma https://orcid.org/0000-0002-7099-7972, Henry Brodaty https://orcid.org/0000-0001-9487-6617, Han G. Brunner, Randy L. Buckner, Jan K. Buitelaar https://orcid.org/0000-0001-8288-7757, Juan R. Bustillo https://orcid.org/0000-0001-8730-8152, Wiepke Cahn https://orcid.org/0000-0002-0482-8759, Murray J. Cairns https://orcid.org/0000-0003-2490-2538, Vince Calhoun https://orcid.org/0000-0001-9058-0747, Vaughan J. Carr https://orcid.org/0000-0002-8907-5804, Xavier Caseras, Svenja Caspers, Gianpiero L. Cavalleri https://orcid.org/0000-0002-9802-0506, Fernando Cendes, Aiden Corvin https://orcid.org/0000-0001-6717-4089, Benedicto Crespo-Facorro https://orcid.org/0000-0003-0033-7132, John C. Dalrymple-Alford https://orcid.org/0000-0002-6672-033X, Udo Dannlowski https://orcid.org/0000-0002-0623-3759, Eco J. C. de Geus https://orcid.org/0000-0001-6022-2666, Ian J. Deary, Norman Delanty, Chantal Depondt https://orcid.org/0000-0002-8452-5319, Sylvane Desrivières https://orcid.org/0000-0002-9120-7060, Gary Donohoe https://orcid.org/0000-0003-3037-7426, Thomas Espeseth https://orcid.org/0000-0001-9076-6365, Guillén Fernández, Simon E. Fisher https://orcid.org/0000-0002-3132-1996, Herta Flor https://orcid.org/0000-0003-4809-5398, Andreas J. Forstner, Clyde Francks https://orcid.org/0000-0002-9098-890X, Barbara Franke https://orcid.org/0000-0003-4375-6572, David C. Glahn, Randy L. Gollub https://orcid.org/0000-0002-9434-4044, Hans J. Grabe, Oliver Gruber, Asta K. Håberg https://orcid.org/0000-0002-9007-1202, Ahmad R. Hariri, Catharina A. Hartman https://orcid.org/0000-0002-8094-8859, Ryota Hashimoto, Andreas Heinz, Frans A. Henskens https://orcid.org/0000-0003-2358-5630, Manon H. J. Hillegers https://orcid.org/0000-0003-4877-282X, Pieter J. Hoekstra https://orcid.org/0000-0003-1018-9954, Avram J. Holmes https://orcid.org/0000-0001-6583-803X, L. Elliot Hong, William D. Hopkins, Hilleke E. Hulshoff Pol https://orcid.org/0000-0002-2038-5281, Terry L. Jernigan, Erik G. Jönsson https://orcid.org/0000-0001-8368-6332, René S. Kahn, Martin A. Kennedy https://orcid.org/0000-0002-6445-8526, Tilo T. J. Kircher https://orcid.org/0000-0002-2514-2625, Peter Kochunov, John B. J. Kwok https://orcid.org/0000-0001-9574-6195, Stephanie Le Hellard https://orcid.org/0000-0002-8085-051X, Carmel M. Loughland, Nicholas G. Martin https://orcid.org/0000-0003-4069-8020, Jean-Luc Martinot, Colm McDonald, Katie L. McMahon https://orcid.org/0000-0002-6357-615X, Andreas Meyer-Lindenberg https://orcid.org/0000-0001-5619-1123, Patricia T. Michie https://orcid.org/0000-0002-4169-8519, Rajendra A. Morey https://orcid.org/0000-0002-6517-6969, Bryan Mowry https://orcid.org/0000-0002-4115-5645, Lars Nyberg, Jaap Oosterlaan https://orcid.org/0000-0002-0218-5630, Roel A. Ophoff https://orcid.org/0000-0002-8287-6457, Christos Pantelis https://orcid.org/0000-0002-9565-0238, Tomas Paus, Zdenka Pausova https://orcid.org/0000-0002-7784-1380, Brenda W. J. H. Penninx https://orcid.org/0000-0001-7779-9672, Tinca J. C. Polderman https://orcid.org/0000-0001-5564-301X, Danielle Posthuma https://orcid.org/0000-0001-7582-2365, Marcella Rietschel https://orcid.org/0000-0002-5236-6149, Joshua L. Roffman https://orcid.org/0000-0003-4073-532X, Laura M. Rowland, Perminder S. Sachdev https://orcid.org/0000-0002-9595-3220, Philipp G. Sämann, Ulrich Schall https://orcid.org/0000-0001-9041-4562, Gunter Schumann, Rodney J. Scott https://orcid.org/0000-0001-7724-3404, Kang Sim https://orcid.org/0000-0003-3209-9626, Sanjay M. Sisodiya, Jordan W. Smoller, Iris E. Sommer https://orcid.org/0000-0001-5597-3262, Beate St Pourcain https://orcid.org/0000-0002-4680-3517, Dan J. Stein https://orcid.org/0000-0001-7218-7810, Arthur W. Toga https://orcid.org/0000-0001-7902-3755, Julian N. Trollor https://orcid.org/0000-0002-7685-2977, Nic J. A. Van der Wee https://orcid.org/0000-0003-3235-1633, Dennis van ’t Ent https://orcid.org/0000-0003-2001-1375, Henry Völzke, Henrik Walter https://orcid.org/0000-0002-9403-6121, Bernd Weber https://orcid.org/0000-0002-7811-9605, Daniel R. Weinberger https://orcid.org/0000-0003-2409-2969, Margaret J. Wright https://orcid.org/0000-0001-7133-4970, Juan Zhou https://orcid.org/0000-0002-0180-8648, Jason L. Stein https://orcid.org/0000-0003-4829-0513 [email protected], Paul M. Thompson https://orcid.org/0000-0002-4720-8867 [email protected], Sarah E. Medland https://orcid.org/0000-0003-1382-380X [email protected], and Enhancing NeuroImaging Genetics through Meta-Analysis Consortium (ENIGMA)—Genetics working groupAuthors Info & Affiliations
Science
20 Mar 2020
Vol 367, Issue 6484

Structured Abstract

INTRODUCTION

The cerebral cortex underlies our complex cognitive capabilities. Variations in human cortical surface area and thickness are associated with neurological, psychological, and behavioral traits and can be measured in vivo by magnetic resonance imaging (MRI). Studies in model organisms have identified genes that influence cortical structure, but little is known about common genetic variants that affect human cortical structure.

RATIONALE

To identify genetic variants associated with human cortical structure at both global and regional levels, we conducted a genome-wide association meta-analysis of brain MRI data from 51,665 individuals across 60 cohorts. We analyzed the surface area and average thickness of the whole cortex and 34 cortical regions with known functional specializations.

RESULTS

We identified 369 nominally genome-wide significant loci (P < 5 × 10−8) associated with cortical structure in a discovery sample of 33,992 participants of European ancestry. Of the 360 loci for which replication data were available, 241 loci influencing surface area and 66 influencing thickness remained significant after replication, with 237 loci passing multiple testing correction (P < 8.3 × 10−10; 187 influencing surface area and 50 influencing thickness).
Common genetic variants explained 34% (SE = 3%) of the variation in total surface area and 26% (SE = 2%) in average thickness; surface area and thickness showed a negative genetic correlation (rG = −0.32, SE = 0.05, P = 6.5 × 10−12), which suggests that genetic influences have opposing effects on surface area and thickness. Bioinformatic analyses showed that total surface area is influenced by genetic variants that alter gene regulatory activity in neural progenitor cells during fetal development. By contrast, average thickness is influenced by active regulatory elements in adult brain samples, which may reflect processes that occur after mid-fetal development, such as myelination, branching, or pruning. When considered together, these results support the radial unit hypothesis that different developmental mechanisms promote surface area expansion and increases in thickness.
To identify specific genetic influences on individual cortical regions, we controlled for global measures (total surface area or average thickness) in the regional analyses. After multiple testing correction, we identified 175 loci that influence regional surface area and 46 that influence regional thickness. Loci that affect regional surface area cluster near genes involved in the Wnt signaling pathway, which is known to influence areal identity.
We observed significant positive genetic correlations and evidence of bidirectional causation of total surface area with both general cognitive functioning and educational attainment. We found additional positive genetic correlations between total surface area and Parkinson’s disease but did not find evidence of causation. Negative genetic correlations were evident between total surface area and insomnia, attention deficit hyperactivity disorder, depressive symptoms, major depressive disorder, and neuroticism.

CONCLUSION

This large-scale collaborative work enhances our understanding of the genetic architecture of the human cerebral cortex and its regional patterning. The highly polygenic architecture of the cortex suggests that distinct genes are involved in the development of specific cortical areas. Moreover, we find evidence that brain structure is a key phenotype along the causal pathway that leads from genetic variation to differences in general cognitive function.
Identifying genetic influences on human cortical structure.
(A) Measurement of cortical surface area and thickness from MRI. (B) Genomic locations of common genetic variants that influence global and regional cortical structure. (C) Our results support the radial unit hypothesis that the expansion of cortical surface area is driven by proliferating neural progenitor cells. (D) Cortical surface area shows genetic correlation with psychiatric and cognitive traits. Error bars indicate SE.
IMAGE CREDITS: (A) K. COURTNEY; (C) M. R. GLASS

Abstract

The cerebral cortex underlies our complex cognitive capabilities, yet little is known about the specific genetic loci that influence human cortical structure. To identify genetic variants that affect cortical structure, we conducted a genome-wide association meta-analysis of brain magnetic resonance imaging data from 51,665 individuals. We analyzed the surface area and average thickness of the whole cortex and 34 regions with known functional specializations. We identified 237 significant loci and found significant enrichment for loci influencing total surface area within regulatory elements that are active during prenatal cortical development, supporting the radial unit hypothesis. Loci that affect regional surface area cluster near genes in Wnt signaling pathways, which influence progenitor expansion and areal identity. Variation in cortical structure is genetically correlated with cognitive function, Parkinson’s disease, insomnia, depression, neuroticism, and attention deficit hyperactivity disorder.

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Supplementary Materials

This PDF file includes:

Materials and Methods
Supplementary Text
Consortium Authors
Additional Cohort Information
Supplementary Acknowledgments
Figs. S1 to S11
Tables S1 to S20
References (70109)

Other Supplementary Material for this manuscript includes the following:

Tables S1 to S20
Figures S2 to S4
Figure S5
Figure S11

References and Notes

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Volume 367 | Issue 6484
20 March 2020

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Acknowledgments

We thank K. Courtney and M. R. Glass for making panels A and C, respectively, of the summary figure. We thank all cohort participants for making this work possible. We thank the research support staff of all cohorts, including interviewers, computer and laboratory technicians, clerical workers, research scientists, volunteers, managers, receptionists, nurses, carers, participating general practitioners, and pharmacists. In addition, the ALSPAC cohort is grateful to the midwives for their help in recruiting the families who participated in the study and to L. B. Clauss for help during the quality control process of the ALSPAC neuroimaging data. The BETULA cohort thanks the Centre for Advanced Study at the Norwegian Academy of Science and Letters in Oslo for hosting collaborative projects and workshops between Norway and Sweden in 2011–2012 and acknowledges that the image analyses were performed on resources provided by the Swedish National Infrastructure for Computing at HPC2N in Umeå. The BONN cohort thanks (in alphabetical order) M. Bartling, U. Broicher, L. Ehrmantraut, A. Maaser, B. Mahlow, S. Mentges, K. Raczka, L. Schinabeck, and P. Trautner for their support and help. The CARDIFF cohort thank researchers at Cardiff University who contributed to the MBBrains panel. The DNS cohort thanks the staff of the Laboratory of NeuroGenetics. The FBIRN cohort thanks L. McMillan for overall study coordination and H. Mangalam, J. Farran, and A. Brenner for administering the University of California, Irvine High-Performance Computing cluster. The GIG cohort thanks M. Keil, E. Diekhof, T. Melcher, and I. Henseler for assistance in data acquisition. The IMpACT cohort acknowledges that, in this work, samples from the Netherlands node of IMpACT were used and the work was carried out on the Dutch national e-infrastructure with the support of the SURF Cooperative. The MCIC cohort thanks colleagues who served as mentors, advisers, and supporters during the inception and conduct of the study, including D. Goff, G. Kuperberg, J. Goldstein, M. Shenton, R. McCarley, S. Heckers, C. Wible, R. Mesholam-Gately, and M. Vangel; staff and clinicians at each site responsible for data acquisition, including S. Wallace, A. Cousins, R. Mesholam-Gately, S. Stufflebeam, O. Freudenreich, D. Holt, L. Kunkel, F. Fleming, G. He, H. Johnson, R. Pierson, A. Caprihan, P. Somers, C. Portal, K. Norman, D. South, M. Doty, and H. Milner; and L. Friedman, S. Posse, J. Jovicich, and T. Wassink for expert guidance on image and other types of data acquisition. MCIC also acknowledges the many research assistants, students, and colleagues who assisted in data curation over the years since data acquisition was completed, including S. Wallace, C. Zyloney, K. Sawlani, J. Fries, A. Scott, D. Wood, R. Wang, W. Courtney, A. Guimaraes, L. Shenkman, M. Kendi, A. T. Karagulle Kendi, R. Muetzel, T. Biehl, and M. Schmidt. The MIRECC cohort thanks the U.S. military veterans who participated in this research. The MPIP cohort thanks R. Schirmer, E. Schreiter, R. Borschke, I. Eidner, and A. Olynyik for supporting MR acquisition and data management, the staff of the Center of Applied Genotyping for generating the genotypes of the Munich Antidepressant Response Signature (MARS) cohort, D. P. Auer for initiating the RUD-MR substudy, E. Binder for supporting participation in ENIGMA, and GlaxoSmithKline for providing the genotypes of the Recurrent Unipolar Depression Case-Control Sample. The PAFIP cohort acknowledges the IDIVAL Neuroimaging Unit for imaging acquirement and analysis and Valdecilla Biobank for help with the technical execution of this work. The PDNZ cohort is grateful to M. MacAskill, D. Myall, L. Livingston, B. Young, and S. Grenfell; staff at the New Zealand Brain Research Institute and Pacific Radiology Christchurch for study coordination and image acquisition; and A. Miller for DNA preparation and banking. The QTIM cohort thanks the many research assistants, radiographers, and IT support staff for data acquisition and DNA sample preparation. The SHIP cohort is grateful to M. Stanke for the opportunity to use his Server Cluster for the SNP imputation, as well as to H. Prokisch and T. Meitinger (Helmholtz Zentrum München) for the genotyping of the SHIP-Trend cohort. The Sydney MAS cohort acknowledges that the genome-wide genotyping was performed by the Ramaciotti Centre, University of New South Wales. Acknowledgments from CHARGE replication cohorts: The Austrian Stroke Prevention Family/Austrian Stroke Prevention Family Study cohort thanks B. Reinhart for her long-term administrative commitment, E. Hofer for technical assistance in creating the DNA bank, I. J. Semmler and A. Harb for DNA sequencing and analyses by TaqMan assays, and I. Poelzl for supervising the quality management processes after ISO9001 at the biobanking and DNA analysis stages. The Cardiovascular Health Study cohort thanks a full list of principal investigators and institutions that can be found at CHS-NHLBI.org. The Framingham Heart Study cohort (CHARGE) especially thanks investigators and staff from the Neurology group for their contributions to data collection. Generation and management of GWAS genotype data for the Rotterdam Study were executed by the Human Genotyping Facility of the Genetic Laboratory of the Department of Internal Medicine, Erasmus MC, Rotterdam, Netherlands. The Rotterdam Study cohort thanks P. Arp, M. Jhamai, M. Verkerk, L. Herrera, M. Peters, and C. Medina-Gomez for help with creating the GWAS database and K. Estrada, Y. Aulchenko, and C. Medina-Gomez for creation and analysis of imputed data. The Three-City Dijon cohort thanks A. Boland (CNG) for technical help in preparing the DNA samples for analyses. The investigators of the frontotemporal GWAS (69), the consortia members, and their acknowledgments are listed in the supplementary materials.
Funding: This study was supported by U54 EB020403 from the National Institutes of Health (NIH) Big Data to Knowledge (BD2K) Initiative, a cross-NIH partnership. Additional support was provided by R01 MH116147, R01 MH1161671, P41 EB015922, RF1 AG051710, RF1 AG041915, R56 AG058854, R01 AG059874, R01 MH117601, the Michael J. Fox Foundation (MJFF; 14848), the Kavli Foundation, and National Health and Medical Research Council (NHMRC) Project Grant 1158127 (to S.E.M.). S.E.M. was funded by an NHMRC Senior Research Fellowship (APP1103623). K.L.G. was supported by APP1173025. L.C.-C. was supported by a QIMR Berghofer Fellowship. J.L.S. was supported by R01MH118349 and R00MH102357. The 1000BRAINS cohort thanks the Heinz Nixdorf Foundation (Germany) for generous support of the Heinz Nixdorf Recall Study, which is also supported by the German Federal Ministry of Education and Science (BMBF; FKZ 01EG940) and the German Research Foundation (DFG; ER 155/6-1). This work was further supported by the BMBF through the Integrated Network IntegraMent under the e:Med Program (01ZX1314A to S.Ci) and the Swiss National Science Foundation (156791 to S.Ci). The Initiative and Networking Fund of the Helmholtz Association supported S.Ca, the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation program (H2020) supported S.Ci (Human Brain Project SGA1, 720270) and supported S.Ca and S.Ci (Human Brain Project SGA2, 785907). The Alzheimer’s Disease Neuroimaging Initiative (ADNI1 and ADNI2GO) was supported by NIH (U01 AG024904) and Department of Defense ADNI (W81XWH-12-2-0012). ADNI is funded by the National Institute on Aging (NIA), the National Institute of Biomedical Imaging and Bioengineering, and through generous contributions from AbbVie; Alzheimer’s Association; Alzheimer’s Drug Discovery Foundation; Araclon Biotech; BioClinica, Inc.; Biogen; Bristol-Myers Squibb Company; CereSpir, Inc.; Cogstate; Eisai Inc.; Elan Pharmaceuticals, Inc.; Eli Lilly and Company; EuroImmun; F. Hoffmann-La Roche, Ltd., and its affiliated company Genentech, Inc.; Fujirebio; GE Healthcare; IXICO Ltd.; Janssen Alzheimer Immunotherapy Research & Development, LLC.; Johnson & Johnson Pharmaceutical Research & Development, LLC; Lumosity; Lundbeck; Merck & Co., Inc.; Meso Scale Diagnostics, LLC; NeuroRx Research; Neurotrack Technologies; Novartis Pharmaceuticals Corporation; Pfizer, Inc.; Piramal Imaging; Servier; Takeda Pharmaceutical Company; and Transition Therapeutics. The Canadian Institutes of Health Research provided funds to support ADNI clinical sites in Canada. Private sector contributions were facilitated by the Foundation for the National Institutes of Health (FNIH) (www.fnih.org). The grantee organization is the Northern California Institute for Research and Education, and the study was coordinated by the Alzheimer’s Therapeutic Research Institute at the University of Southern California. Samples used in this study were from the National Centralized Repository for Alzheimer’s Disease and Related Dementias, which received government support under a cooperative agreement grant (U24 AG21886) awarded by the NIA. Support for data analysis was provided by NLM R01 LM012535 and NIA R03 AG054936 (to K.N.). The UK Medical Research Council (MRC) and Wellcome (102215/2/13/2) and the University of Bristol provide core support for ALSPAC. A comprehensive list of grants funding is available on the ALSPAC website (www.bristol.ac.uk/alspac/external/documents/grant-acknowledgements.pdf). ALSPAC neuroimaging data were specifically funded by RO1 MH085772 (to T.P.). GWAS data were generated by Sample Logistics and Genotyping Facilities at Wellcome Sanger Institute and LabCorp (Laboratory Corporation of America), with support from 23andMe. Data and sample collection by the Australian Schizophrenia Research Bank (ASRB) were supported by the Australian NHMRC, the Pratt Foundation, Ramsay Health Care, and the Viertel Charitable Foundation. The ASRB was also supported by the Schizophrenia Research Institute (Australia), with infrastructure funding from NSW Health and the Macquarie Group Foundation. DNA analysis was supported by the Neurobehavioral Genetics Unit, with funding from NSW Health and the NHMRC Project Grants (1067137, 1147644, 1051672). M.C. was supported by a NHMRC Senior Research Fellowship (1121474). C.P. was supported by a NHMRC Senior Principal Research Fellowship (628386 and 1105825). BETULA was supported by a Wallenberg Scholar grant from the Knut and Alice Wallenberg Foundation and a grant from Torsten and Ragnar Söderbergs Foundation to L.N., HelseVest RHF (911554 to S.L.H.), grants from the Bergen Research Foundation and the University of Bergen to S.L.H., grants from the Dr Einar Martens Fund and the K.G. Jebsen Foundation to S.L.H. and V.M.S., and the Research Council of Norway (177458/V 50 to T.E. and 204966/F 20 to L.T.W.). Nijmegen’s BIG resource is part of Cognomics, a joint initiative by researchers of the Donders Centre for Cognitive Neuroimaging, the Human Genetics and Cognitive Neuroscience departments of the Radboud University Medical Center, and the Max Planck Institute for Psycholinguistics (funded by the Max Planck Society). Support for the Cognomics Initiative, including phenotyping and genotyping of BIG cohorts, comes from funds of the participating departments and centers and from external national grants: the Biobanking and Biomolecular Resources Research Infrastructure (Netherlands) (BBMRI-NL), the Hersenstichting Nederland, and the Netherlands Organisation for Scientific Research (NWO), including the NWO Brain & Cognition Excellence Program (433-09-229) and the Vici Innovation Program (016-130-669 to B.F.). Additional support was received from the European Union’s Seventh Framework Program (FP7) [602805 (Aggressotype), 602450 (IMAGEMEND), and 278948 (TACTICS)], from H2020 [643051 (MiND) and 667302 (CoCA)], and from the Innovative Medicines Initiative 2 Joint Undertaking (H2020/EFPIA) [115916 (PRISM)]. BONN was supported by the Frankfurt Institute for Risk Management and Regulation, and B.W. was supported by a Heisenberg Grant of the DFG [WE 4427 (3-2)]. BrainScale was supported by NWO (NWO 51.02.061 to H.E.H.P., NWO 51.02.062 to D.I.B., NWO-NIHC Programs of excellence 433-09-220 to H.E.H.P., NWO-MagW 480-04-004 to D.I.B., and NWO/SPI 56-464-14192 to D.I.B.), FP7 Ideas: European Research Council (ERC-230374 to D.I.B.), and Universiteit Utrecht (High Potential Grant to H.E.H.P.). CARDIFF genotyping was supported by the National Centre for Mental Health. T.M.L. is funded by a Sêr Cyrmu II Fellowship (East Wales European Regional Development Funds (PNU-80762-CU-14) at the Dementia Research Institute, Cardiff University. DNS received support from Duke University as well as NIH (R01DA033369 and R01DA031579). Work from the London cohort of EPIGEN was supported by research grants from the Wellcome Trust (084730 to S.M.S.), University College London (UCL)/University College London Hospitals (UCLH) National Institute for Health Research (NIHR) Biomedical Research Centre/Specialist Biomedical Research Centres (CBRC/SBRC) (114 to S.M.S.), the Comprehensive Local Research Network Flexibility and Sustainability Funding (CEL1300 to S.M.S.), The Big Lottery Fund, the Wolfson Trust, and the Epilepsy Society. This work was partly undertaken at UCLH/UCL, which received a proportion of funding from the NIHR CBRC/SBRC. FBIRN was supported by the NIH National Center for Research Resources (NCRR) [NIH 1 U24 RR021992 (Function Biomedical Informatics Research Network), NIH 1 U24 RR025736-01 (Biomedical Informatics Research Network Coordinating Center)], the NIH National Center for Research Resources and the National Center for Advancing Translational Sciences (UL1 TR000153), and the NIH through 5R01MH094524, and P20GM103472. This work was supported in part by a Merit Review Award I01CX000497 (J.M.F.) and a Senior Research Career Award (J.M.F.) from the U.S. Department of Veterans Affairs, Clinical Sciences Research and Development Service. FOR2107 was funded by the DFG (FOR2107 DA1151/5-1, DA1151/5-2 to U.D.; JA1890/7-1, JA1890/7-2 to A.J.; KI 588/14-1, KI 588/14-2 to T.K.; KR 3822/7-1, KR 3822/7-2 to A.K.; NO246/10-1, NO246/10-2 to M.M.N.). GOBS was supported by the National Institute of Mental Health MH0708143 (to D.C.G.), MH078111 (to J.Bl.), and MH083824 (to D.C.G. and J.Bl.). The Brain Genomics Superstruct Project (GSP) was made possible by the resources provided through Shared Instrumentation Grants 1S10RR023043 and 1S10RR023401 and was supported by funding from the Simons Foundation (to R.L.B.); the Howard Hughes Medical Institute (to R.L.B.); NIMH grants R01-MH079799 (to J.W.S.), K24MH094614 (to J.W.S.), and K01MH099232 (to A.J.H.); and the Massachusetts General Hospital–University of Southern California Human Connectome Project (U54MH091665). The HUBIN cohort was supported by the Swedish Research Council (2006-2992, 2006-986, K2007-62X-15077-04-1, K2008-62P-20597-01-3, 2008-2167, 2008-7573, K2010-62X-15078-07-2, K2012-61X-15078-09-3, 14266-01A, 02-03, 2017-949), the regional agreement on medical training and clinical research between Stockholm County Council and the Karolinska Institutet, the Knut and Alice Wallenberg Foundation, and the HUBIN project. HUNT-MRI was funded by the Liaison Committee between the Central Norway Regional Health Authority and the Norwegian University of Science and Technology, and the Norwegian National Advisory Unit for functional MRI. IMAGEN was supported by the European Union’s FP6 Integrated Project IMAGEN (LSHM-CT- 2007-037286), the H2020 ERC Advanced Grant STRATIFY (695313), ERANID (PR-ST-0416-10004), BRIDGET (JPND: MR/N027558/1), the FP7 projects IMAGEMEND (602450) and MATRICS (603016), the Innovative Medicine Initiative Project EU-AIMS (115300-2), the Medical Research Foundation and MRC (MR/R00465X/1), the MRC (MR/N000390/1), the Swedish Research Council FORMAS, the NIHR Biomedical Research Centre at South London and Maudsley NHS Foundation Trust and King’s College London, the BMBF (01GS08152, 01EV0711, eMED SysAlc01ZX1311A, Forschungsnetz AERIAL), and the DFG (SM 80/7-1, SM 80/7-2, SFB 940/1). Further support was provided by ANR (AF12-NEUR0008-01 – WM2NA, and ANR-12-SAMA-0004), the Fondation de France, the Fondation pour la Recherche Médicale, the Mission Interministérielle de Lutte-contre-les-Drogues-et-les-Conduites-Addictives (MILDECA), the Fondation pour la Recherche Médicale (DPA20140629802), the Fondation de l’Avenir, Paris Sud University IDEX 2012, Science Foundation Ireland (16/ERCD/3797), and the NIH (RO1 MH085772-01A1). IMH was supported by National Healthcare Group, Singapore (SIG/05004; SIG/05028), and Singapore Bioimaging Consortium (RP C-009/2006) research grants awarded to K.S.; M.La. was supported by a National Medical Research Council Research Training Fellowship (MH095: 003/008-1014) and Singapore Ministry of Health National Medical Research Council Center Grant (NMRC/CG/004/2013). IMpACT was supported by the NWO (433-09-229) and the Vici Innovation Program (016-130-669 to B.F.). Additional support was received from the ERC under FP7 [602805 (Aggressotype), 602450 (IMAGEMEND), and 278948 (TACTICS)] as well as from H2020 [643051 (MiND), 667302 (CoCA), and 728018 (Eat2beNICE)]. LBC1936 was supported by a Research into Aging program grant (to I.J.D.) and the Age UK-funded Disconnected Mind project www.disconnectedmind.ed.ac.uk; to I.J.D. and J.M.W.), with additional funding from the UK MRC (Mr/M01311/1, G1001245/96077, G0701120/79365 to I.J.D., J.M.W. and M.E.B.). The whole-genome association part of this study was funded by the Biotechnology and Biological Sciences Research Council (BBSRC; BB/F019394/1). J.M.W. is supported by the Scottish Funding Council through the SINAPSE Collaboration (www.sinapse.ac.uk). CCACE (MRC MR/K026992/1) is funded by the BBSRC and MRC. LIBD was supported by direct funding from the NIMH intramural research program of the NIH to the Weinberger laboratory and by support from the Lieber Institute for Brain Development and the Maltz Research Laboratories. MCIC was supported primarily by U.S. Department of Energy DE-FG02-99ER62764 through its support of the Mind Research Network (MRN, formerly known as the MIND Institute) and the consortium as well as by the National Association for Research in Schizophrenia and Affective Disorders (NARSAD) Young Investigator Award (to S.Eh.), through the Blowitz-Ridgeway and Essel Foundations and a ZonMw TOP 91211021 (to T.Wh.), a DFG research fellowship (to S.Eh.), the MRN, the NIH through NCRR 5MO1-RR001066 (MGH General Clinical Research Center), NIMH K08 MH068540, the Biomedical Informatics Research Network with NCRR Supplements to P41 RR14075 (MGH), M01 RR 01066 (MGH), NIBIB R01EB006841 (MRN), R01EB005846 (MRN), 2R01 EB000840 (MRN), 1RC1MH089257 (MRN), as well as grant U24 RR021992. Meth-CT was supported by the Medical Research Council, South Africa. MIRECC was supported by NIMH (1R01MH111671) and the U.S. Department of Veterans Affairs (VISN6 MIRECC). MooDS was supported by the BMBF grants [National Genome Research Network Plus (MooDS: Systematic Investigation of the Molecular Causes of Major Mood Disorders and Schizophrenia, www.ngfn.de/en/schizophrenie.html); e:Med Programme: Integrated Network IntegraMent (01ZX1314A to M.M.N.; 01ZX1614A to F.D. and M.M.N.)] and by DFG (FOR 1617) as well as Excellence Cluster (EXC 257). MPIP was supported by a grant of the Exzellenz-Stiftung of the Max Planck Society and by the BMBF National Genome Research Network (FKZ 01GS0481). MPRC was supported by the NIH (R01MH116948, R01MH112180, U01MH108148, UG3DA047685, 2R01EB015611, R01DA027680, R01MH085646, P50MH103222, U54 EB020403, and T32MH067533), NSF (IIS-1302755 and MRI-1531491), a state of Maryland contract (M00B6400091), and a Pfizer research grant. MÜNSTER was funded by the DFG (SFB-TRR58, Projects C09 and Z02 to U.D.) and the Interdisciplinary Center for Clinical Research of the medical faculty of Münster (Dan3/012/17 to U.D.). NCNG was supported by the Bergen Research Foundation, the University of Bergen, the Research Council of Norway [FUGE (151904 and 183327), Psykisk Helse (175345), RCN (154313/V50 to I.R. and 177458/V50 to T.E.)], Helse Sørøst RHF (2012086 to T.E.), and the Dr Einar Martens Fund. NESDA obtained funding from the NWO (Geestkracht program 10-000-1002), the Center for Medical Systems Biology (CSMB, NWO Genomics), BBMRI-NL, VU University’s Institutes for Health and Care Research (EMGO+) and Neuroscience Campus Amsterdam, University Medical Center Groningen, Leiden University Medical Center, and NIH (R01D0042157-01A, MH081802, Grand Opportunity grants 1RC2 MH089951 and 1RC2 MH089995). Parts of the genotyping and analyses efforts were funded by the Genetic Association Information Network (GAIN) of the FNIH. Computing was supported by BiG Grid, the Dutch e-Science Grid, which is financially supported by NWO. The NeuroIMAGE study was supported by NIH grant R01MH62873, NWO Large Investment Grant 1750102007010 (to J.K.B.), ZonMW grant 60-60600-97-193, NWO grants 056-13-015 and 433-09-242, and matching grants from Radboud University Nijmegen Medical Center, University Medical Center Groningen and Accare, and Vrije Universiteit Amsterdam. Further support was received from the FP7 [278948 (TACTICS), 602450 (IMAGEMEND), 602805 (Aggressotype)], and H2020 [667302 (CoCA) and 728018 (Eat2beNICE)]. The Netherlands Twin Register (NTR) obtained funding from NWO and ZonMW grants (904-61-090, 985-10-002, 912-10-020, 904-61-193,480-04-004, 463-06-001, 451-04-034, 400-05-717, Addiction-31160008, 016-115-035, 481-08-011, 056-32-010, Middelgroot-911-09-032, OCW_NWO Gravity program –024.001.003, NWO-Groot 480-15-001/674), Center for Medical Systems Biology (CSMB, NWO Genomics), NBIC/BioAssist/RK (2008.024), BBMRI–NL (184.021.007 and 184.033.111); Spinozapremie (NWO- 56-464-14192), KNAW Academy Professor Award (PAH/6635), and University Research Fellow grant to D.I.B.; Amsterdam Public Health research institute (former EMGO+), Neuroscience Amsterdam research institute (former NCA); the European Science Foundation (EU/QLRT-2001-01254), FP7 [FP7- HEALTH-F4-2007-2013: 01413 (ENGAGE) and 602768 (ACTION)]; the ERC (ERC Advanced, 230374, ERC Starting grant 284167), Rutgers University Cell and DNA Repository (NIMH U24 MH068457-06), the NIH (R01D0042157-01A1, R01MH58799-03, MH081802, DA018673, R01 DK092127-04, Grand Opportunity grants 1RC2 MH089951, and 1RC2 MH089995); and the Avera Institute for Human Genetics. Parts of the genotyping and analyses efforts were funded by the Genetic Association Information Network (GAIN) of the FNIH. Computing was supported by NWO through 2018/EW/00408559, BiG Grid, the Dutch e-Science Grid and SURFSARA. OATS was supported by the Australian NHMRC/Australian Research Council Strategic Award (401162) and NHMRC Project Grant (1405325). The study was facilitated through Twins Research Australia, a national resource partly supported by a Centre for Research Excellence from the NHMRC. DNA was extracted by Genetic Repositories Australia (NHMRC grant 401184). Genome-wide genotyping at the Diamantina Institute, University of Queensland, was partly funded by a CSIRO Flagship Collaboration Fund Grant. OSAKA was supported by AMED under JP18dm0307002, JP18dm0207006 (Brain/MINDS) and JSPS KAKENHI J16H05375. PAFIP was supported by the Instituto de Salud Carlos III (PI14/00639 and PI14/00918), MINECO (SAF2010-20840-C02-02 and SAF2013-46292-R), and Fundación Instituto de Investigación Marqués de Valdecilla (NCT0235832 and NCT02534363). PDNZ was supported by the Health Research Council, the Neurological Foundation of New Zealand, Canterbury Medical Research Foundation, University of Otago Research Grant, and Jim and Mary Carney Charitable Trust (Whangarei, New Zealand). PING was supported by the National Institute on Drug Abuse (RC2DA029475) and the U.S. National Institute of Child Health and Human Development (R01HD061414). Parkinson’s Progression Markers Initiative (PPMI), a public-private partnership, is funded by the Michael J. Fox Foundation for Parkinson’s Research and funding partners, including AbbVie, Allegran, Avid Radiopharmaceuticals, Biogen Idec, BioLegend, Bristol-Meyers Squibb, Denali Therapeutics, GE Healthcare, Genentech, GSK-GlaxoSmithKline, Eli Lilly & Co., F. Hoffman-La Roche Ltd., Lundbeck Pharmaceuticals, Merck & Co., MSD-Meso Scale Discovery, Pfizer, Piramal, Sanofi Genzyme, Servier, Takeda Pharmaceutical Company, TEVA Pharmaceutical Industries, UCB Pharma SA, and Golub Capital (www.ppmi-info.org/about-ppmi/who-we-are/study-sponsors/). QTIM was supported by the National Institute of Child Health and Human Development (R01 HD050735), National Institute of Biomedical Imaging and Bioengineering (Award 1U54EB020403-01, Subaward 56929223), and NHMRC (Project Grants 496682, 1009064 and Medical Bioinformatics Genomics Proteomics Program 389891). SHIP is part of the Community Medicine Research net of the University of Greifswald, Germany, which is funded by the DFG (01ZZ9603, 01ZZ0103, and 01ZZ0403), the Ministry of Cultural Affairs, the Social Ministry of the Federal State of Mecklenburg-West Pomerania, and the network “Greifswald Approach to Individualized Medicine (GANI_MED)” funded by the DFG (03IS2061A). Whole-body MRI was supported by a joint grant from Siemens Healthineers, Erlangen, Germany, and the federal state of Mecklenburg–West Pomerania. Genome-wide data have been supported by the DFO (03ZIK012) and a joint grant from Siemens Healthineers, Erlangen, Germany, and the federal state of Mecklenburg–West Pomerania. The University of Greifswald is a member of the Caché Campus program of InterSystems, GmbH. Sydney MAS was supported by the NHMRC/Australian Research Council Strategic Award (401162) and NHMRC Program Grants (350833, 568969). DNA was extracted by Genetic Repositories Australia (NHMRC Grant 401184). SYS has been funded by the Canadian Institutes of Health Research and the Heart and Stroke Foundation of Canada. Computations were performed on the GPC supercomputer at the SciNet HPC Consortium. SciNet is funded by the Canada Foundation for Innovation under the auspices of Compute Canada; the government of Ontario; Ontario Research Fund–Research Excellence, and the University of Toronto. TCD-NUIG included data from two sites. NUI Galway data collection was supported by the Health Research Board (HRA_POR/2011/100). Trinity College Dublin was supported by The Science Foundation Ireland Research Investigator project (12.IP.1359 to G.D.). TOP and TOP3T are part of TOP, which is supported by the Research Council of Norway (223273, 213837, 249711, 226971, 262656), the South East Norway Health Authority (2017-112), the Kristian Gerhard Jebsen Stiftelsen (SKGJ-MED-008), and the FP7 [602450 (IMAGEMEND)]. UiO2016 and UiO2017 are part of TOP and STROKEMRI, which are supported by the Norwegian ExtraFoundation for Health and Rehabilitation (2015/FO5146), the Research Council of Norway (249795, 248238), and the South-Eastern Norway Regional Health Authority (2014097, 2015044, 2015073). The UMCU cohort consists of several independent studies, which were supported by ZonMw TOP 40-008-12-98-13009, the Geestkracht program of the ZonMw (10-000-1001), the Stanley Medical Research Institute (Dr. Nolen), the Brain and Behavior Research Foundation (2013-2015 NARSAD Independent Investigator grant 20244 to M.H.J.H.), NWO (2012-2017 VIDI grants 452-11-014 to N.E.M.v.H. and 917-46-370 to H.E.H.P.), and ZonMw (908-02-123 to H.E.H.P.). UNICAMP was supported by FAPESP (São Paulo Research Foundation) 2013/07559-3: the Brazilian Institute of Neuroscience and Neurotechnology (BRAINN). Collection of chimpanzee brain images was supported by the NIH (NS-42867, NS-73134, and NS-92988). The views expressed in this manuscript are those of the authors and do not necessarily represent the views of the National Heart, Lung, and Blood Institute; the NIH; or the U.S. Department of Health and Human Services. Infrastructure for the CHARGE Consortium is supported in part by the National Heart, Lung, and Blood Institute (NHLBI; HL105756) and for the neuroCHARGE phenotype working group through the NIA (AG033193). H.H.H.A. was supported by the Netherlands Organization for the Health Research and Development (ZonMw; 916.19.151). The Atherosclerosis Risk in Communities Study (ARIC) was a collaborative study supported by the NHLBI (HHSN268201100005C, HSN268201100006C, HSN268201100007C, HHSN268201100008C, HHSN268201100009C, HHSN268201100010C, HHSN268201100011C, HHSN268201100012C, R01HL70825, R01HL087641, R01HL59367, and R01HL086694); the National Human Genome Research Institute (U01HG004402); and the NIH (HHSN268200625226C). Infrastructure was partly supported by UL1RR025005, a component of the NIH and NIH Roadmap for Medical Research. This project was partially supported by NIH R01 grants HL084099 and NS087541 (to M.Fo.). The Austrian Stroke Prevention Family/Austrian Stroke Prevention Family Study databank was supported by the Medical University of Graz and the Steiermärkische Krankenanstaltengesellschaft. The research reported in this article was funded by the Austrian Science Fund (PI904, P20545-P05 and P13180), the Austrian National Bank Anniversary Fund (P15435), and the Austrian Ministry of Science under the aegis of the EU Joint Programme–Neurodegenerative Disease Research (JPND; www.jpnd.eu). The Cardiovascular Health Study was supported by NHLBI (HHSN268201200036C, HHSN268200800007C, N01HC55222, N01HC85079, N01HC85080, N01HC85081, N01HC85082, N01HC85083, N01HC85086, U01HL080295, R01HL087652, R01HL105756, R01HL103612, R01HL120393, and R01HL130114), with additional contribution from the National Institute of Neurological Disorders and Stroke. Additional support was provided through R01AG023629, R01AG15928, and R01AG033193 from the NIA. The provision of genotyping data was supported in part by the National Center for Advancing Translational Sciences, CTSI grant UL1TR000124, and the National Institute of Diabetes and Digestive and Kidney Disease Diabetes Research Center grant DK063491 to the Southern California Diabetes Endocrinology Research Center. The Erasmus Rucphen Family Study was supported by the Consortium for Systems Biology, within the framework of the Netherlands Genomics Initiative (NGI)/NWO. ERF as a part of EUROSPAN (European Special Populations Research Network) was supported by the European Commission’s 5th Framework Programme (FP5) (QLG2-CT-2002-01254), the FP6 (018947; LSHG-CT-2006-01947), and the FP7 (HEALTH-F4-2007-201413 and 602633). High-throughput analysis of the ERF data was supported by a joint grant from the Netherlands Organisation for Scientific Research and the Russian Foundation for Basic Research (NWO-RFBR 047.017.043). High-throughput metabolomics measurements of the ERF study were supported by BBMRI-NL. The Framingham Heart Study was supported by the National Heart, Lung, and Blood Institute’s Framingham Heart Study (N01-HC-25195 and HHSN268201500001I) and its contract with Affymetrix, Inc., for genotyping services (N02-HL-6-4278). A portion of this research used the Linux Cluster for Genetic Analysis (LinGA-II) funded by the Robert Dawson Evans Endowment of the Department of Medicine at Boston University School of Medicine and Boston Medical Center. This study was also supported by grants from the NIA (R01s AG033040, AG033193, AG054076, AG049607, AG008122, AG016495, and U01-AG049505) and the National Institute of Neurological Disorders and Stroke (R01-NS017950). C.Dec. is supported by the Alzheimer’s Disease Center (P30 AG 010129). LIFE-Adult: LIFE-Adult is funded by the Leipzig Research Center for Civilization Diseases (LIFE). LIFE is an organizational unit affiliated with the Medical Faculty of the University of Leipzig. LIFE is funded by means of the European Union, the European Regional Development Fund, and funds of the Free State of Saxony within the framework of the excellence initiative. This work was also funded by the DFG (CRC 1052 “Obesity mechanisms” project A1) and the Max Planck Society. The Rotterdam Study was funded by Erasmus Medical Center and Erasmus University, Rotterdam; ZonMw; the Research Institute for Diseases in the Elderly (RIDE); the Ministry of Education, Culture and Science; the Ministry for Health, Welfare and Sports; the European Commission (DG XII); and the municipality of Rotterdam. The GWAS datasets are supported by NWO Investments (175.010.2005.011, 911-03-012); the Genetic Laboratory of the Department of Internal Medicine, Erasmus MC; the Research Institute for Diseases in the Elderly (014-93-015; RIDE2); and the NGI/NWO Netherlands Consortium for Healthy Aging (050-060-810). This work was performed as part of the CoSTREAM project (www.costream.eu) and received funding from H2020 (667375). The Three-City Dijon study was conducted under a partnership agreement among the Institut National de la Santé et de la Recherche Médicale (INSERM), the University of Bordeaux, and Sanofi-Aventis. The Fondation pour la Recherche Médicale funded the preparation and initiation of the study. The 3C Study is also supported by the Caisse Nationale Maladie des Travailleurs Salariés, Direction Générale de la Santé, Mutuelle Générale de l’Education Nationale, Institut de la Longévité, Conseils Régionaux of Aquitaine and Bourgogne, Fondation de France, and Ministry of Research–INSERM Programme “Cohortes et collections de données biologiques.” S.Deb. received investigator-initiated research funding from the French National Research Agency (ANR) and from the Fondation Leducq. S.Deb. is supported by a starting grant from the European Research Council (SEGWAY), a grant from the Joint Programme of Neurodegenerative Disease research (BRIDGET), H2020 (643417 and 640643), and the Initiative of Excellence of Bordeaux University. This work was supported by the National Foundation for Alzheimer’s Disease and Related Disorders, the Institut Pasteur de Lille, the labex DISTALZ, and the Centre National de Génotypage. The Vietnam Era Twin Study of Aging (VETSA) was supported by the U.S. NIH VA San Diego Center of Excellence for Stress and Mental Health R00DA023549, DA-18673, NIA R01 AG018384, R01 AG018386, R01 AG022381, R01 AG022982, R01 DA025109 05, R01 HD050735, K08 AG047903, R03 AG 046413, and R01 HD050735-01A2.
Author contributions: Central analysis and coordination group: C.R.K.C., D.P.H., F.P., J.Br., J.L.S., J.N.P., K.L.G., L.C.-C., L.C.P.Z., L.T.S., M.A.B.M., N.J., N.S., P.A.L., P.M.T., S.E.M., and S.I.T. Manuscript writing, preparation, and revision: A.H.Z., D.P.H., J.Br., J.L.S., J.N.P., K.L.G., L.C.-C., N.J., P.A.L., P.M.T., and S.E.M. Project support: D.Ga., M.A.B.M., M.J., N.S., R.E., V.R., and Y.G. Cohort principal investigators: A.A.V., A.C., A.H., A.J.F., A.J.H., A.K.H., A.M.D., A.M.-L., A.R.H., A.W.T., B.C.-F., B.F., B.Mo., B.S.P., B.W., B.W.J.H.P., C.A.H., C.Dep., C.F., C.M., C.M.L., C.P., D.Am., D.C.G., D.I.B., D.J.S., D.P., D.R.W., D.v.E., E.G.J., E.J.C.d.G., L.E.H., F.A.H., F.C., G.D., G.F., G.G.B., G.L.C., G.S., H.B., H.E.H.P., H.F., H.G.B., H.J.G., H.V., H.W., I.A., I.E.Som., I.J.D., I.M., J.B.J.K., J.Bl., J.C.Be., J.C.D.-A., J.K.B., J.-L.M., J.L.R., J.N.T., J.O., J.R.B., J.W.S., J.Z., K.L.M., K.S., L.M.R., L.N., L.R., L.T.W., M.E.B., M.H.J.H., M.J.C., M.J.W., M.K.M.A., M.R., N.D., N.J., N.J.A.v.d.W., O.A.A., O.G., P.G.S., P.J.H., P.K., P.M.T., P.S.S., P.T.M., R.A.M., R.A.O., R.H., R.J.S., R.L.B., R.L.G., R.S.K., S.Ca., S.Des., S.E.F., S.L.H., S.M.S., S.R., T.E., T.J.A., T.J.C.P., T.L.J., T.P., T.T.J.K., U.D., V.C., V.J.C., W.C., W.U.H., X.C., and Z.P. Imaging data collection: A.B., A.d.B., A.F.M., A.J., A.J.H., A.K., A.K.H., A.L.G., A.M.D., A.N.H., A.P., A.R.H., A.R.K., A.U., B.A.M., B.-C.H., B.D., B.F., G.B.P., B.W., B.W.J.H.P., C.B., C.D.W., C.J., C.L.B., C.L.Y., C.M., C.P., C.R.J., C.S.Re., D.Am., D.C.G., D.Gr., D.H.M., D.J., D.J.H., D.J.V., D.M.C., D.P.O., D.R.W., D.S.O., D.T.-G., D.v.E., D.v.R., D.Z., E.A., E.B.Q., E.J.C.d.G., L.E.H., E.Sh., G.B.P., G.D., G.F., G.I.d.Z., G.L.C., G.R., G.S., H.V., H.Y., I.A., I.E.Som., J.A.T., J.E.C., J.E.N., J.K.B., J.-L.M., J.-L.M., J.L.R., J.M.F., J.M.W., J.N.T., J.R., J.T.V., K.D., K.K., K.L.M., K.O.L., K.S., L.M.R., L.R., L.T.W., M.B.H., M.E.B., M.Fu., M.H.J.H., M.Ho., M.-J.v.T., M.J.W., M.-L.P.M., N.E.M.v.H., N.F.H., N.H., N.J.A.v.d.W., N.K.H., N.O., O.G., P.A.G., P.E.R., P.G.S., P.K., P.N., P.S.S., R.A.O., R.B., R.H., R.L.B., R.L.G., R.R., R.S.K., R.W., S.A., S.C.M., S.Ca., S.Er., S.Ko., S.M., S.M.S., T.G.M.v.E., T.R.M., T.Wh., T.W.M., U.D., U.S., V.C., V.J.C., V.S.M., W.D.H., W.H., W.W., and X.C. Imaging data analysis: A.F.M., A.H.Z., A.J.H., A.J.S., A.L.G., A.M.D., A.R., A.R.K., A.S., A.Th., A.U., B.A.G., B.C.R., B.F., B.K., B.S.P., C.B., C.C.F., C.C.H., C.D.W., C.J., C.L.Y., C.R.K.C., C.S.Ro., D.Al., D.C.G., D.Gr., D.H., D.J., D.J.H., D.M.C., D.P.H., D.P.O., D.T.-G., D.v.d.M., D.v.E., D.v.R., D.Z., E.E.L.B., E.Sh., E.Sp., E.W., F.M.R., F.P., F.S., G.I.d.Z., G.R., H.J.G., I.A., I.E.Som., I.K.A., J.A.T., J.B.J.K., J.C.V.M., J.-L.M., J.L.R., J.L.S., J.M.W., J.R., J.Z., K.D., K.L.M., K.N., K.S., K.W., L.B.L., L.H., L.Sa., L.Sc., L.Sh., L.T.S., L.T.W., L.v.E., L.C.P.Z., M.A., M.A.H., M.B.H., M.C., M.E.B., M.Fu., M.Ho., M.J.G., M.-J.v.T., M.J.W., M.Ki., M.La., M.P.Z., M.W., N.E.M.v.H., N.F.H., N.J., N.O., N.T.D., O.G., P.G.S., P.K., P.M.T., P.N., R.B., R.K., R.L.G., R.M.B., R.R., S.A., S.Ca., S.Des., S.Eh., S.Er., S.F.F., S.I.T., S.Ka., S.Ke., S.L.R., S.M.C.d.Z., S.R.M., T.A., T.A.L., T.G., T.G.M.v.E., T.J., T.K., T.L.P., T.P.G., T.R.M., T.Wh., T.Wo., T.W.M., U.D., W.W., X.C., Y.Q., and Z.Z. Genetic data collection: A.A.A., A.A.-K., A.d.B., A.J.F., A.J.H., A.J.S., A.K.H., A.M.D., A.P., A.R.H., A.R.K., B.-C.H., B.F., B.Mo., B.T.B., B.W., B.W.J.H.P., C.B., C.D.W., C.F., C.M., C.P., C.P.D., C.S.Re., D.C.G., D.H.M., D.R.W., D.W.M., D.Z., E.A., E.B.Q., E.G.J., E.J.C.d.G., L.E.H., F.D., F.M., F.R.T., G.D., G.E.D., G.F., G.H., G.L.C., G.S., H.V., H.Y., I.E.Som., I.L.-C., J.A.T., J.B.J.K., J.Bl., J.E.C., J.E.N., J.-J.H., J.J.L., J.K.B., J.-L.M., J.-L.M., J.L.R., J.M.F., J.Q.W., J.R., J.W.S., K.A.M., K.D., K.O.L., K.S., L.M.R., L.R., L.Sh., M.A.K., M.F.D., M.H.J.H., M.Ha., M.Ho., M.J.C., M.J.W., M.La., M.-L.P.M., M.M.N., M.N., N.A.K., N.E.M.v.H., N.G.M., N.J.A.v.d.W., N.K.H., N.O., O.G., P.A.T., P.H., P.K., P.R.S., P.S.S., R.A.O., R.C.G., R.H., R.L.B., R.R., R.Se., R.S.K., R.W., S.A., S.Ci., S.Dj., S.E.F., S.Eh., S.Er., S.H., S.L.H., S.M.S., T.G.M.v.E., T.J.A., T.K.d.A., T.L.P., T.W.M., U.D., V.C., V.J.C., V.M.S., and X.C. Genetic data analysis: A.A.-K., A.J.F., A.J.H., A.J.S., A.M.D., A.R.K., A.Te., A.Th., B.C.-D., B.F., B.K., B.M.-M., B.P., B.S.P., B.T.B., C.C.F., C.D.W., C.L.V., C.S.Re., C.S.Ro., C.W., C.Y.S., D.C.G., D.K., D.P.H., D.v.d.M., D.v.E., E.G.J., L.E.H., E.V., E.W., F.M., H.-R.E., I.E.J., I.E.Som., I.E.Søn., I.L.-C., I.O.F., J.Bl., J.Br., J.F.P., J.H.V., J.-J.H., J.L.R., J.L.S., J.N.P., J.Q.W., J.R.A., J.S., J.W.C., J.W.S., K.E.T., K.L.G., K.N., L.C.-C., L.M.O.L., L.Sh., L.C.P.Z., M.A.A.A., M.B., M.E.G., M.Fu., M.Ha., M.I., M.J., M.J.C., M.J.W., M.Ki., M.Kl., M.Kn., M.La., M.Lu., M.M.J.v.D., N.A.G., N.G.M., N.J., N.J.A., N.K.H., N.M.-S., N.R.M., O.G., P.A.L., P.G.S., P.H., P.H.L., P.K., P.M.T., P.R.S., Q.C., R.A.O., R.M.B., R.R., R.Se., S.Da., S.Des., S.E.M., S.Eh., S.G., S.H., S.H.W., S.L.H., S.M.C.d.Z., S.N., S.R.M., T.A.L., T.G., T.G.M.v.E., T.J., T.K.d.A., T.M.L., W.R.R., Y.M., and Y.W. CHARGE study design: B.Ma., C.Dec., C.L.S., E.H., G.V.R., H.H.H.A., H.J.G., J.C.Bi., L.L., M.A.I., M.Fo., O.L.L., Q.Y., R.Sc., S.Deb., S.S., T.H.M., T.P., V.G., and W.T.L.
Competing interests: A.M.D. is a Founder of CorTechs Labs, Inc., and has received funding through a Research Agreement between General Electric Healthcare and the University of California, San Diego. The terms of this arrangement have been reviewed and approved by the University of California, San Diego, in accordance with its conflict of interest policies. B.F. has received educational speaking fees from Shire and Medice. B.W.J.H.P. has received (nonrelated) research funding from Boehringer Ingelheim and Janssen Research. C.D.W. is currently an employee of Biogen. C.R.J. consults for Lilly and serves on an independent data monitoring board for Roche but receives no personal compensation from any commercial entity. C.R.J. receives research support from NIH and the Alexander Family Alzheimer’s Disease Research Professorship of the Mayo Clinic. D.H.M. is a consultant for Boehringer Ingelheim, GW/Greenwhich Biosciences, and Aptinyx. D.J.S. has received research grants and/or consultancy honoraria from Lundbeck and Sun. D.P.H. is currently an employee of Genentech, Inc., and was previously employed by Janssen R&D, LLC. H.B. is on the advisory board for Nutricia Australia. R.S.K. has consulted for Alkermes, Otsuka, Luye Pharma, and Sunovion and has received speaker fees from Janssen-Cilag and Lundbeck. None of the other authors declare any competing financial interests. H.J.G. has received travel grants and speaker honoraria from Fresenius Medical Care, Neuraxpharm, and Janssen Cilag. H.J.G. has received research funding from the German Research Foundation, the German Ministry of Education and Research, the DAMP Foundation, Fresenius Medical Care, the EU Joint Programme Neurodegenerative Disorders, and the European Social Fund. L.E.H. has received or is planning to receive research funding or consulting fees from Mitsubishi; Your Energy Systems, LLC; Neuralstem; Taisho; Heptares; Pfizer; Sound Pharma; Luye Pharma; Takeda; and Regeneron. N.H. is a stockholder of Siemens AG, Erlangen, Germany. R.B. has received travel grants and speaker honoraria from Bayer Healthcare, AG. R.L.B. is a paid consultant for Roche.
Data and materials availability: The meta-analytic results presented in this paper are available to download from the ENIGMA consortium website (http://enigma.ini.usc.edu/research/download-enigma-gwas-results). Access to cohort data is available either through public repositories or directly from the cohort. Direct requests are required when informed consent or the approved study protocol does not permit deposition into a repository. Requests for data by qualified investigators are subject to scientific and ethical review to ensure that the data will be used for valid scientific research and to ensure compliance with confidentiality, consent, and data protection regulations. Some of the data are subject to material transfer agreements or data transfer agreements, and specific details on how to access data for each cohort are available in table S20.

Authors

Affiliations

Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
School of Psychology, University of Queensland, Brisbane, QLD, Australia.
School of Biomedical Sciences, Queensland University of Technology, Brisbane, QLD, Australia.
Faculty of Psychology, University of Murcia, Murcia, Spain.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Derrek P. Hibar
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Personalized Healthcare, Genentech, Inc., South San Francisco, CA, USA.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
School of Biomedical Sciences, Queensland University of Technology, Brisbane, QLD, Australia.
Faculty of Medicine, University of Queensland, Brisbane, QLD, Australia.
Roles: Formal analysis, Investigation, Validation, Writing - original draft, and Writing - review & editing.
Fabrizio Pizzagalli
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Formal analysis, Software, Validation, Visualization, and Writing - review & editing.
Christopher R. K. Ching
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Graduate Interdepartmental Program in Neuroscience, University of California Los Angeles, Los Angeles, CA, USA.
Roles: Conceptualization, Data curation, Formal analysis, Methodology, Project administration, Software, Supervision, and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Investigation, Project administration, Writing - original draft, and Writing - review & editing.
Natalia Shatokhina
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Role: Software.
Department of Genetics and UNC Neuroscience Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
Roles: Data curation, Formal analysis, Software, and Validation.
Sophia I. Thomopoulos
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Role: Investigation.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Visualization and Writing - review & editing.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Roles: Formal analysis and Investigation.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Department of Psychiatric Research, Diakonhjemmet Hospital, Oslo, Norway.
Centre for Psychiatric Research, Department of Clinical Neuroscience, Karolinska Institutet, Stockholm, Sweden.
Roles: Conceptualization, Funding acquisition, Investigation, Project administration, and Resources.
Saud Alhusaini
Department of Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.
Neurology Department, Yale School of Medicine, New Haven, CT, USA.
Roles: Data curation, Formal analysis, Investigation, and Resources.
Department of Human Genetics and South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine, Brownsville, TX, USA.
Roles: Data curation, Formal analysis, Investigation, Methodology, Software, and Writing - review & editing.
Dag Alnæs
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Data curation and Formal analysis.
Centre for Lifespan Changes in Brain and Cognition, Department of Psychology, University of Oslo, Oslo, Norway.
Roles: Formal analysis and Resources.
Department of Integrative Medical Biology, Umeå University, Umeå, Sweden.
Umeå Center for Functional Brain Imaging, Umeå University, Umeå, Sweden.
Roles: Data curation and Formal analysis.
Laboratory of Neuro Imaging, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of the University of Southern California, Los Angeles, CA, USA.
Role: Software.
Mathematics and Statistics, Murdoch University, Murdoch, WA, Australia.
Roles: Formal analysis and Writing - review & editing.
Allison Ashley-Koch
Duke Molecular Physiology Institute, Duke University Medical Center, Durham, NC, USA.
Roles: Formal analysis, Supervision, and Writing - review & editing.
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.
Roles: Formal analysis, Investigation, Methodology, and Software.
The Hospital for Sick Children, University of Toronto, Toronto, ON, Canada.
Roles: Data curation, Formal analysis, Methodology, Resources, and Software.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Data curation, Investigation, Resources, and Writing - review & editing.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Role: Formal analysis.
Robin Bülow
Institute for Radiology and Neuroradiology, University Medicine, Ernst-Moritz-Arndt University, Greifswald, Germany.
Roles: Conceptualization, Data curation, Investigation, Project administration, Resources, and Validation.
Christian Bürger
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Data curation, Investigation, and Validation.
Dara M. Cannon
Centre for Neuroimaging and Cognitive Genomics, National University of Ireland Galway, Galway, Ireland.
Roles: Funding acquisition, Investigation, Project administration, Resources, Supervision, and Writing - review & editing.
Mallar Chakravarty
Douglas Mental Health University Institute, McGill University, Montreal, QC, Canada.
Departments of Psychiatry and Biological and Biomedical Engineering, McGill University, Montreal, QC, Canada.
Roles: Data curation, Investigation, Validation, and Writing - review & editing.
Lieber Institute for Brain Development, Baltimore, MD, USA.
Roles: Formal analysis and Investigation.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Formal analysis, Investigation, and Software.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Institute for Molecular Bioscience, University of Queensland, Brisbane, QLD, Australia.
Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
Roles: Data curation, Formal analysis, and Software.
Department of Neurosciences, University of California, San Diego, La Jolla, CA, USA.
Department of Radiology, University of California San Diego, San Diego, CA, USA.
Roles: Conceptualization, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, and Supervision.
Shareefa Dalvie
Department of Psychiatry and Mental Health, University of Cape Town, Cape Town, South Africa.
Roles: Formal analysis and Resources.
Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Roles: Investigation and Resources.
Faculty of Health, Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, QLD, Australia.
Roles: Conceptualization, Funding acquisition, Investigation, Resources, and Supervision.
Sonja M. C. de Zwarte
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Formal analysis, Investigation, and Resources.
Anouk den Braber
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Vrije Universiteit Amsterdam, Amsterdam UMC, Amsterdam, Netherlands.
Roles: Formal analysis and Investigation.
Nhat Trung Doan
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Role: Formal analysis.
Department of Psychiatry, University of Münster, Münster, Germany.
Role: Investigation.
Division of Psychological and Social Medicine and Developmental Neurosciences, Faculty of Medicine, Technische Universität Dresden, Dresden, Germany.
Roles: Formal analysis, Investigation, and Resources.
Division of Human Genetics, Institute of Infectious Disease and Molecular Medicine, University of Cape Town, Cape Town, South Africa.
Roles: Formal analysis and Software.
Susanne Erk
Division of Mind and Brain Research, Department of Psychiatry and Psychotherapy, Campus Charité Mitte, Charité - Universitätsmedizin Berlin corporate member of Freie Universität Berlin, Humboldt-Universität zu Berlin, and Berlin Institute of Health, Berlin, Germany.
Roles: Formal analysis and Investigation.
Chun Chieh Fan
Department of Cognitive Science, University of California San Diego, San Diego, CA, USA.
Roles: Formal analysis, Investigation, Software, and Validation.
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Role: Formal analysis.
Sonya F. Foley
Cardiff University Brain Research Imaging Centre, Cardiff University, Cardiff, UK.
Role: Investigation.
San Francisco Veterans Administration Medical Center, San Francisco, CA, USA.
Roles: Data curation, Funding acquisition, Investigation, Resources, Supervision, and Validation.
Division of Cerebral Integration, National Institute for Physiological Sciences, Okazaki, Japan.
Roles: Formal analysis, Investigation, Resources, Software, and Validation.
Duke Molecular Physiology Institute, Duke University Medical Center, Durham, NC, USA.
Role: Formal analysis.
Tian Ge
Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA.
Athinoula A. Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Boston, MA, USA.
Roles: Data curation, Formal analysis, Investigation, and Software.
Sudheer Giddaluru
NORMENT - K.G. Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway.
Roles: Formal analysis, Investigation, Validation, Writing - original draft, and Writing - review & editing.
Aaron L. Goldman
Lieber Institute for Brain Development, Baltimore, MD, USA.
Roles: Formal analysis and Investigation.
School of Psychiatry, University of New South Wales, Sydney, NSW, Australia.
Neuroscience Research Australia, Sydney, NSW, Australia.
Roles: Investigation, Resources, and Writing - review & editing.
Nynke A. Groenewold
Department of Psychiatry and Mental Health, University of Cape Town, Cape Town, South Africa.
Roles: Data curation and Formal analysis.
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Data curation, Formal analysis, Investigation, Resources, and Software.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Department of Psychiatric Research, Diakonhjemmet Hospital, Oslo, Norway.
Roles: Formal analysis and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Department of Biomedical Engineering, Illinois Institute of Technology, Chicago, IL, USA.
Role: Software.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Roles: Data curation and Writing - review & editing.
Centre for Clinical Brain Sciences and Edinburgh Imaging, University of Edinburgh, Edinburgh, UK.
Division of Psychiatry, University of Edinburgh, Edinburgh, UK.
Roles: Conceptualization, Formal analysis, and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Formal analysis, Project administration, Software, and Validation.
Duke UNC Brain Imaging and Analysis Center, Duke University Medical Center, Durham, NC, USA.
Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham VA Medical Center, Durham, NC, USA.
Roles: Formal analysis and Investigation.
Duke Molecular Physiology Institute, Duke University Medical Center, Durham, NC, USA.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.
Department of Genomics, Life & Brain Research Center, University of Bonn, Bonn, Germany.
Roles: Data curation and Resources.
Dirk J. Heslenfeld
Department of Cognitive and Clinical Neuropsychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Funding acquisition, Investigation, Project administration, Resources, and Writing - review & editing.
New Fei Ho
Research Division, Institute of Mental Health, Singapore, Singapore.
Roles: Investigation and Resources.
Max Planck Institute of Psychiatry, Munich, Germany.
Roles: Investigation and Resources.
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.
Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany.
Roles: Investigation and Resources.
Centre for Neuroimaging and Cognitive Genomics, School of Psychology, National University of Ireland Galway, Galway, Ireland.
Roles: Conceptualization, Formal analysis, Validation, and Writing - review & editing.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Roles: Data curation, Formal analysis, and Validation.
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Data curation, Formal analysis, Investigation, Resources, Software, and Validation.
Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Japan.
Roles: Formal analysis, Investigation, and Resources.
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany.
Roles: Conceptualization, Data curation, Investigation, Methodology, Resources, Validation, Visualization, and Writing - review & editing.
Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Department of Neurology, Alzheimer Center, Amsterdam Neuroscience, Vrije Universiteit Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Role: Formal analysis.
Institute of Science and Technology for Brain-Inspired Intelligence, Fudan University, Shanghai, China.
Key Laboratory of Computational Neuroscience and Brain-Inspired Intelligence (Fudan University), Ministry of Education, Shanghai, China.
Centre for Population Neuroscience and Precision Medicine (PONS), Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK.
Roles: Formal analysis and Software.
Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.
Department of Psychiatry, Psychotherapy and Psychosomatics, Medical Faculty, RWTH Aachen University, Aachen, Germany.
JARA-BRAIN, Jülich-Aachen Research Alliance, Jülich, Germany.
Roles: Formal analysis, Investigation, and Resources.
Department of Neuroinformatics, Araya, Inc., Tokyo, Japan.
Sackler Centre for Consciousness Science, School of Psychology, University of Sussex, Falmer, UK.
Earth-Life Science Institute, Tokyo Institute of Technology, Tokyo, Japan.
Roles: Formal analysis and Software.
Douglas Mental Health University Institute, McGill University, Montreal, QC, Canada.
Department of Psychiatry, McGill University, Montreal, QC, Canada.
McConnell Brain Imaging Center, Montreal Neurological Institute, Montreal, QC, Canada.
Roles: Formal analysis, Investigation, and Resources.
Dalia Kasperaviciute
Department of Clinical and Experimental Epilepsy, UCL Queen Square Institute of Neurology, London, UK.
Genomics England, Queen Mary University of London, London, UK.
Roles: Formal analysis, Investigation, Software, Validation, and Writing - review & editing.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Data curation and Formal analysis.
Sinead Kelly
Public Psychiatry Division, Massachusetts Mental Health Center, Beth Israel Deaconess Medical Center, Harvard Medical School, Boston, MA, USA.
Psychiatry Neuroimaging Laboratory, Department of Psychiatry, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.
Roles: Data curation, Formal analysis, Investigation, Resources, Software, Validation, and Writing - review & editing.
Department of Genome Informatics, Graduate School of Medicine, Osaka University, Suita, Japan.
Role: Formal analysis.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Formal analysis, Investigation, Software, and Validation.
Michael Knapp
Department of Medical Biometry, Informatics and Epidemiology, University Hospital Bonn, Bonn, Germany.
Roles: Formal analysis, Resources, and Software.
Department of Psychology and Neuroscience, Duke University, Durham, NC, USA.
Roles: Data curation, Formal analysis, and Investigation.
Section for Experimental Psychopathology and Neuroimaging, Department of General Psychiatry, Heidelberg University Hospital, Heidelberg, Germany.
Centre for Translational Research in Systems Neuroscience and Psychiatry, Department of Psychiatry and Psychotherapy, University Medical Center Göttingen, Göttingen, Germany.
Roles: Data curation, Formal analysis, Investigation, and Validation.
Max Lam
Research Division, Institute of Mental Health, Singapore, Singapore.
Human Genetics, Genome Institute of Singapore, Singapore, Singapore.
Roles: Formal analysis, Investigation, Resources, Software, and Writing - review & editing.
Thomas M. Lancaster
Cardiff University Brain Research Imaging Centre, Cardiff University, Cardiff, UK.
MRC Centre for Neuropsychiatric Genetics and Genomics, Cardiff University, Cardiff, UK.
Roles: Data curation, Formal analysis, Investigation, and Software.
Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA.
Department of Psychiatry, Harvard Medical School, Boston, MA, USA.
Roles: Formal analysis, Investigation, Resources, Software, and Writing - review & editing.
Tristram A. Lett
Division of Mind and Brain Research, Department of Psychiatry and Psychotherapy, Campus Charité Mitte, Charité - Universitätsmedizin Berlin corporate member of Freie Universität Berlin, Humboldt-Universität zu Berlin, and Berlin Institute of Health, Berlin, Germany.
Roles: Formal analysis and Investigation.
McConnell Brain Imaging Center, Montreal Neurological Institute, Montreal, QC, Canada.
McGill Centre for Integrative Neuroscience, McGill University, Montreal, QC, Canada.
Roles: Formal analysis and Resources.
Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Department of Psychology, University of Edinburgh, Edinburgh, UK.
Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.
Role: Formal analysis.
Department of Psychiatry and Human Behavior, School of Medicine University of California, Irvine, Irvine, CA, USA.
Roles: Conceptualization, Resources, and Validation.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.
Roles: Conceptualization, Data curation, Formal analysis, Methodology, Validation, and Writing - review & editing.
Samuel R. Mathias
Department of Psychiatry, Yale University School of Medicine, New Haven, CT, USA.
Olin Neuropsychiatric Research Center, Institute of Living, Hartford Hospital, Hartford, CT, USA.
Roles: Investigation and Resources.
Department of Medicine, University of Otago, Christchurch, Christchurch, New Zealand.
New Zealand Brain Research Institute, Christchurch, New Zealand.
Brain Research New Zealand - Rangahau Roro Aotearoa, Christchurch, New Zealand.
Roles: Investigation, Resources, and Writing - review & editing.
Yuri Milaneschi
Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, Netherlands.
Roles: Formal analysis and Writing - review & editing.
Max Planck Institute of Psychiatry, Munich, Germany.
Institute of Neurogenomics, Helmholtz Zentrum München, German Research Centre for Environmental Health, Neuherberg, Germany.
Role: Formal analysis.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
IC - Institute of Computing, Campinas, Brazil.
Roles: Formal analysis, Investigation, and Resources.
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.
Cécile and Oskar Vogt Institute of Brain Research, Medical Faculty, Heinrich Heine University, Düsseldorf, Germany.
Roles: Formal analysis, Investigation, and Resources.
Bertram Müller-Myhsok
Max Planck Institute of Psychiatry, Munich, Germany.
Munich Cluster for Systems Neurology (SyNergy), Munich, Germany.
Institute of Translational Medicine, Liverpool, UK.
Roles: Formal analysis, Resources, Software, Supervision, and Writing - review & editing.
Centre for Neuroimaging and Cognitive Genomics, National University of Ireland Galway, Galway, Ireland.
Roles: Data curation, Formal analysis, Investigation, Resources, and Validation.
Soichiro Nakahara
Department of Psychiatry and Human Behavior, School of Medicine University of California, Irvine, Irvine, CA, USA.
Drug Discovery Research, Astellas Pharmaceuticals, 21 Miyukigaoka, Tsukuba, Ibaraki 305-8585, Japan.
Role: Formal analysis.
Kwangsik Nho
Department of Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, IN, USA.
Roles: Formal analysis, Investigation, Software, and Writing - review & editing.
Center for Neurobehavioral Genetics, University of California Los Angeles, Los Angeles, CA, USA.
Role: Formal analysis.
Dimitri Papadopoulos Orfanos https://orcid.org/0000-0002-1242-8990
NeuroSpin, CEA, Université Paris-Saclay, Gif-sur-Yvette, France.
Roles: Data curation, Investigation, Methodology, Resources, and Software.
Biostatistics and Computational Biology Unit, University of Otago, Christchurch, Christchurch, New Zealand.
Department of Pathology and Biomedical Science, University of Otago, Christchurch, Christchurch, New Zealand.
Roles: Data curation, Formal analysis, Software, and Validation.
Department of Medicine, University of Otago, Christchurch, Christchurch, New Zealand.
New Zealand Brain Research Institute, Christchurch, New Zealand.
Brain Research New Zealand - Rangahau Roro Aotearoa, Christchurch, New Zealand.
Roles: Investigation and Resources.
Benno Pütz
Max Planck Institute of Psychiatry, Munich, Germany.
Roles: Resources and Writing - review & editing.
School of Psychiatry, University of New South Wales, Sydney, NSW, Australia.
Neuroscience Research Australia, Sydney, NSW, Australia.
Roles: Formal analysis, Investigation, Resources, and Writing - review & editing.
Anjanibhargavi Ragothaman https://orcid.org/0000-0001-9243-9368
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Formal analysis, and Software.
Faisal M. Rashid
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation, Formal analysis, Methodology, Resources, Software, and Validation.
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.
Roles: Formal analysis and Software.
Ronny Redlich
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Resources, Validation, and Writing - review & editing.
Centre for Lifespan Changes in Brain and Cognition, Department of Psychology, University of Oslo, Oslo, Norway.
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.
Roles: Formal analysis, Investigation, and Writing - review & editing.
Department of Psychiatry, University of Münster, Münster, Germany.
Role: Resources.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Department of Psychology, University of Oslo, Oslo, Norway.
Sunnaas Rehabilitation Hospital HT, Nesodden, Norway.
Roles: Investigation and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Department of Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, IN, USA.
Roles: Conceptualization, Formal analysis, Investigation, and Validation.
Shannon L. Risacher
Department of Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, IN, USA.
Roles: Formal analysis, Software, and Validation.
Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Roles: Formal analysis, Resources, and Software.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Psychiatry, Radboud university medical center, Nijmegen, Netherlands.
Roles: Formal analysis and Validation.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Formal analysis, Investigation, Methodology, Resources, and Validation.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Methodology and Software.
Department of Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, IN, USA.
Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, IN, USA.
Roles: Formal analysis, Investigation, Project administration, Resources, and Writing - review & editing.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Roles: Formal analysis, Investigation, Resources, and Writing - review & editing.
Lianne Schmaal
Orygen, The National Centre of Excellence for Youth Mental Health, Melbourne, VIC, Australia.
The Centre for Youth Mental Health, University of Melbourne, Melbourne, VIC, Australia.
Department of Psychiatry, Vrije Universiteit University Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Conceptualization, Formal analysis, Investigation, Resources, and Writing - review & editing.
Neuroscience Research Australia, Sydney, NSW, Australia.
School of Medical Sciences, University of New South Wales, Sydney, NSW, Australia.
Roles: Funding acquisition, Investigation, and Resources.
Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Roles: Formal analysis, Investigation, Resources, and Writing - review & editing.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Roles: Data curation, Formal analysis, Investigation, and Resources.
Department of Biostatistics, Epidemiology and Informatics, University of Pennsylvania, Philadelphia, PA, USA.
Roles: Data curation, Formal analysis, and Investigation.
The Hospital for Sick Children, University of Toronto, Toronto, ON, Canada.
Population Neuroscience & Developmental Neuroimaging, Bloorview Research Institute, University of Toronto, East York, ON, Canada.
Role: Formal analysis.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Donders Centre for Cognitive Neuroimaging, Radboud University, Nijmegen, Netherlands.
Roles: Data curation, Formal analysis, and Investigation.
Ida E. Sønderby
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Formal analysis and Writing - review & editing.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Roles: Formal analysis, Software, and Writing - review & editing.
Katherine E. Tansey
MRC Centre for Neuropsychiatric Genetics and Genomics, Cardiff University, Cardiff, UK.
Roles: Formal analysis, Investigation, Software, Validation, Writing - original draft, and Writing - review & editing.
Alexander Teumer
Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.
Roles: Formal analysis, Investigation, Software, and Writing - review & editing.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Roles: Formal analysis, Investigation, and Software.
Diana Tordesillas-Gutiérrez
Neuroimaging Unit, Technological Facilities, Valdecilla Biomedical Research Institute IDIVAL, Santander, Spain.
Centro Investigacion Biomedica en Red Salud Mental, Santander, Spain.
Department of Psychology, Georgia State University, Atlanta, GA, USA.
Mind Research Network, Albuquerque, NM, USA.
Roles: Conceptualization, Data curation, Investigation, Methodology, Resources, Validation, and Writing - review & editing.
Anne Uhlmann
Department of Psychiatry and Mental Health, University of Cape Town, Cape Town, South Africa.
Department of Psychiatry, University of Vermont, Burlington, VT, USA.
Roles: Investigation and Writing - review & editing.
Costanza Ludovica Vallerga https://orcid.org/0000-0001-9493-6800
Institute for Molecular Bioscience, University of Queensland, Brisbane, QLD, Australia.
Roles: Formal analysis and Resources.
Dennis van der Meer
NORMENT, Division of Mental Health and Addiction, Oslo University Hospital and Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
School of Mental Health and Neuroscience, Faculty of Health, Medicine and Life Sciences, Maastricht University, Maastricht, Netherlands.
Roles: Formal analysis and Investigation.
Marjolein M. J. van Donkelaar
Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Role: Formal analysis.
School of Psychology, University of Queensland, Brisbane, QLD, Australia.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Roles: Formal analysis and Investigation.
Department of Psychiatry and Human Behavior, School of Medicine University of California, Irvine, Irvine, CA, USA.
Roles: Data curation, Formal analysis, Investigation, Resources, and Writing - review & editing.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Department of Child and Adolescent Psychiatry/Psychology, Erasmus Medical Center-Sophia Children’s Hospital, Rotterdam, Netherlands.
Roles: Data curation, Investigation, Resources, and Writing - review & editing.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.
Roles: Investigation and Writing - review & editing.
Cognitive Neuroscience Center, Department of Biomedical Sciences of Cells and Systems, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.
Roles: Data curation, Investigation, Resources, and Writing - review & editing.
Department of Neurology, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Resources and Writing - review & editing.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Roles: Data curation, Formal analysis, Investigation, and Writing - review & editing.
Department of Psychology, Georgia State University, Atlanta, GA, USA.
MRC Integrative Epidemiology Unit, Department of Population Health Sciences, Bristol Medical School, Bristol, UK.
Department of Psychology, University of Bath, Bath, UK.
Roles: Data curation, Formal analysis, Investigation, Software, Validation, and Writing - review & editing.
Research Division, Institute of Mental Health, Singapore, Singapore.
Role: Investigation.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Formal analysis, Investigation, Resources, and Validation.
Centre for Clinical Brain Sciences and Edinburgh Imaging, University of Edinburgh, Edinburgh, UK.
Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.
UK Dementia Research Institute, The University of Edinburgh, Edinburgh, UK.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Resources, Software, Validation, and Writing - review & editing.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Roles: Data curation, Investigation, and Resources.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Department of Psychology, University of Oslo, Oslo, Norway.
Roles: Conceptualization, Formal analysis, Investigation, Methodology, and Writing - review & editing.
Christopher D. Whelan
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Department of Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.
Roles: Data curation, Formal analysis, Investigation, Resources, Software, Validation, and Writing - review & editing.
Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.
Roles: Investigation, Supervision, and Validation.
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany.
German Center for Neurodegenerative Diseases Rostock/Greifswald, Greifswald, Germany.
Roles: Formal analysis and Writing - review & editing.
Department of Psychiatry, Psychosomatics and Psychotherapy, University of Würzburg, Würzburg, Germany.
Roles: Formal analysis, Investigation, Methodology, Software, Validation, Visualization, and Writing - review & editing.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Role: Investigation.
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Resources, Software, Validation, and Writing - review & editing.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Department of Neurology, FCM, UNICAMP, Campinas, Brazil.
Roles: Formal analysis, Investigation, and Resources.
Dario Zaremba
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Formal analysis, Investigation, Validation, and Writing - review & editing.
Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK.
Role: Formal analysis.
Marcel P. Zwiers
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.
Donders Centre for Cognitive Neuroimaging, Radboud University, Nijmegen, Netherlands.
Roles: Data curation, Formal analysis, Investigation, Validation, and Writing - review & editing.
INSERM Unit 1000 - Neuroimaging & Psychiatry, Paris Saclay University, Gif sur Yvette, France.
Role: Investigation.
Amelia A. Assareh
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Roles: Investigation and Resources.
Centro Investigacion Biomedica en Red Salud Mental, Santander, Spain.
Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria–IDIVAL, Santander, Spain.
Roles: Project administration and Resources.
Duke UNC Brain Imaging and Analysis Center, Duke University Medical Center, Durham, NC, USA.
Department of Psychiatry and Frank Porter Graham Child Development Institute, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
Roles: Conceptualization, Investigation, Methodology, Project administration, Resources, Validation, and Writing - review & editing.
Christine L. Brandt
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Investigation and Resources.
Gregory G. Brown
Department of Psychiatry, University of California San Diego, San Diego, CA, USA.
VA San Diego Healthcare System, San Diego, CA, USA.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Project administration, and Resources.
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.
Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.
Roles: Funding acquisition, Investigation, Resources, and Writing - review & editing.
Department of Human Genetics and South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine, Brownsville, TX, USA.
Roles: Data curation, Investigation, and Resources.
Gareth E. Davies
Avera Institute for Human Genetics, Sioux Falls, SD, USA.
Roles: Formal analysis, Investigation, Project administration, Resources, Supervision, and Validation.
Franziska Degenhardt
Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany.
Roles: Investigation and Resources.
Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham VA Medical Center, Durham, NC, USA.
Roles: Investigation, Project administration, Resources, and Validation.
Bruno Dietsche
Department of Psychiatry and Psychotherapy, Philipps-University Marburg, Marburg, Germany.
Role: Investigation.
Department of Medical Genetics, Oslo University Hospital, Oslo, Norway.
NORMENT, Department of Clinical Science, University of Bergen, Bergen, Norway.
Roles: Conceptualization, Data curation, Funding acquisition, Project administration, Resources, Validation, and Writing - review & editing.
Department of Neurology, St James’s Hospital, Dublin, Ireland.
Academic Unit of Neurology, TBSI, Dublin, Ireland.
Future Neuro, Royal College of Surgeons in Ireland, Dublin, Ireland.
Roles: Investigation and Writing - review & editing.
Ryan Espiritu
Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation and Software.
Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.
Roles: Data curation and Software.
Yolanda Gil
Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.
Role: Software.
Sir Peter Mansfield Imaging Centre, University of Nottingham, Nottingham, UK.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, and Validation.
Brigham and Women’s Hospital, Boston, MA, USA.
The Broad Institute, Boston, MA, USA.
Harvard Medical School, Boston, MA, USA.
Roles: Funding acquisition, Project administration, Resources, and Writing - review & editing.
Center for Economics and Neuroscience, University of Bonn, Bonn, Germany.
Institute of Experimental Epileptology and Cognition Research, University Hospital Bonn, Germany.
Roles: Formal analysis and Investigation.
Walter Heindel
Department of Clinical Radiology, University of Münster, Münster, Germany.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, and Writing - review & editing.
Department of Psychiatry, University of Iowa College of Medicine, Iowa City, IA, USA.
Roles: Data curation, Investigation, and Resources.
Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.
German Center for Neurodegenerative Diseases Rostock/Greifswald, Greifswald, Germany.
Roles: Funding acquisition, Investigation, and Writing - review & editing.
Max Planck Institute of Psychiatry, Munich, Germany.
HMNC Holding GmbH, Munich, Germany.
Roles: Conceptualization, Funding acquisition, Project administration, Resources, Supervision, and Writing - review & editing.
University Medicine Greifswald, Interfaculty Institute for Genetics and Functional Genomics, Department of Functional Genomics, Greifswald, Germany.
Roles: Investigation and Resources.
Institute of Diagnostic Radiology and Neuroradiology, Greifswald, Germany.
Roles: Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, and Writing - review & editing.
Department of Radiology, Mayo Clinic, Rochester, MN, USA.
Roles: Data curation, Resources, and Writing - review & editing.
Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.
Roles: Formal analysis, Methodology, and Software.
Department of Psychiatry and Psychotherapy, Philipps-University Marburg, Marburg, Germany.
Core-Unit Brainimaging, Faculty of Medicine, University of Marburg, Marburg, Germany.
Roles: Funding acquisition and Resources.
Nathan A. Kimbrel
Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham VA Medical Center, Durham, NC, USA.
Department of Psychiatry and Behavioral Sciences, Duke University School of Medicine, Durham, NC, USA.
Roles: Investigation, Resources, and Writing - review & editing.
Knut Kolskår
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Department of Psychology, University of Oslo, Oslo, Norway.
Sunnaas Rehabilitation Hospital HT, Nesodden, Norway.
Role: Investigation.
Sanne Koops
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Investigation and Writing - review & editing.
Department of Psychiatry and Psychotherapy, Philipps-University Marburg, Marburg, Germany.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Software, and Writing - review & editing.
Department of Psychiatry, University of Minnesota, Minneapolis, MN, USA.
Roles: Data curation, Investigation, Methodology, Project administration, Resources, Validation, and Writing - review & editing.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Department of Translational Neuroscience, UMC Utrecht Brain Center, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
GGNet Mental Health, Apeldoorn, Netherlands.
Roles: Formal analysis and Writing - review & editing.
Department of Psychiatry and Weill Institute for Neurosciences, University of California San Francisco, San Francisco, CA, USA.
Mental Health Service 116d, Veterans Affairs San Francisco Healthcare System, San Francisco, CA, USA.
Roles: Funding acquisition, Investigation, Project administration, Supervision, and Writing - review & editing.
Karen A. Mather
Neuroscience Research Australia, Sydney, NSW, Australia.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Roles: Investigation, Resources, and Writing - review & editing.
Lieber Institute for Brain Development, Baltimore, MD, USA.
Department of Neurology, Johns Hopkins University, Baltimore, MD, USA.
Department of Radiology, Johns Hopkins University, Baltimore, MD, USA.
Roles: Investigation and Resources.
Sarah Matthews
MRC Integrative Epidemiology Unit, Department of Population Health Sciences, Bristol Medical School, Bristol, UK.
Roles: Data curation, Software, and Validation.
Jaqueline Mayoral Van Son
Centro Investigacion Biomedica en Red Salud Mental, Santander, Spain.
Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria–IDIVAL, Santander, Spain.
Roles: Conceptualization, Investigation, Resources, Validation, and Writing - review & editing.
Pacific Brain Health Center, Santa Monica, CA, USA.
John Wayne Cancer Institute, Santa Monica, CA, USA.
Roles: Data curation, Investigation, Project administration, and Resources.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Investigation, Project administration, Resources, and Writing - review & editing.
Centre for Neuroimaging and Cognitive Genomics, National University of Ireland Galway, Galway, Ireland.
Roles: Investigation and Resources.
Department of Psychiatry, University of Minnesota, Minneapolis, MN, USA.
Roles: Investigation and Writing - review & editing.
Matthias Nauck
Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.
German Centre for Cardiovascular Research, Greifswald, Germany.
Roles: Investigation and Resources.
Sunnaas Rehabilitation Hospital HT, Nesodden, Norway.
Roles: Investigation and Resources.
Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany.
Roles: Funding acquisition, Investigation, Project administration, and Resources.
Department of Psychiatry, University of Iowa College of Medicine, Iowa City, IA, USA.
Roles: Conceptualization and Investigation.
Nils Opel
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Conceptualization and Writing - review & editing.
Marie-Laure Paillère Martinot
INSERM Unit 1000 - Neuroimaging & Psychiatry, Paris Saclay University, Gif sur Yvette, France.
Child and Adolescent Psychiatry Department, APHP Pitié Salpêtrière Hospital, Paris, France.
Roles: Funding acquisition, Investigation, and Resources.
G. Bruce Pike
Radiology and Clinical Neurosciences, Hotchkiss Brain Institute, University of Calgary, Calgary, AB, Canada.
Roles: Investigation, Methodology, Project administration, and Resources.
School of Medicine, University of California Irvine, Irvine, CA, USA.
Roles: Investigation, Validation, and Writing - review & editing.
Erin B. Quinlan
Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK.
Role: Investigation.
Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.
School of Medicine and Public Health, University of Newcastle, Callaghan, NSW, Australia.
Priority Centre for Stroke and Brain Injury, University of Newcastle, Callaghan, NSW, Australia.
Hunter Medical Research Institute, Newcastle, NSW, Australia.
Roles: Data curation, Investigation, Resources, and Writing - review & editing.
Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.
Role: Software.
Simone Reppermund
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Department of Developmental Disability Neuropsychiatry, University of New South Wales, Sydney, NSW, Australia.
Role: Investigation.
NORMENT, Department of Clinical Science, University of Bergen, Bergen, Norway.
Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway.
Role: Resources.
Paul A. Tooney
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Hunter Medical Research Institute, Newcastle, NSW, Australia.
Role: Resources.
Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Role: Investigation.
Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, Netherlands.
Roles: Data curation, Formal analysis, Investigation, Resources, and Writing - review & editing.
Duke UNC Brain Imaging and Analysis Center, Duke University Medical Center, Durham, NC, USA.
Role: Investigation.
School of Psychology, Trinity College Dublin, Dublin, Ireland.
Roles: Funding acquisition, Investigation, Project administration, Resources, and Supervision.
Tonya White
Department of Child and Adolescent Psychiatry/Psychology, Erasmus Medical Center-Sophia Children’s Hospital, Rotterdam, Netherlands.
Department of Radiology, Erasmus University Medical Centre, Rotterdam, Netherlands.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Validation, and Writing - review & editing.
Department of Psychiatry, Osaka University Graduate School of Medicine, Suita, Japan.
Roles: Investigation, Resources, and Writing - review & editing.
Department of Epidemiology, Erasmus MC Medical Center, Rotterdam, Netherlands.
Department of Radiology and Nuclear Medicine, Erasmus MC Medical Center, Rotterdam, Netherlands.
Department of Clinical Genetics, Erasmus MC Medical Center, Rotterdam, Netherlands.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, and Writing - review & editing.
Joshua C. Bis
Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA.
Roles: Formal analysis, Investigation, and Software.
Stephanie Debette
INSERM, Bordeaux Population Health Research Center, team VINTAGE, UMR 1219, University of Bordeaux, Bordeaux, France.
Department of Neurology, CHU de Bordeaux, Bordeaux, France.
Roles: Investigation, Resources, Validation, and Writing - review & editing.
Charles Decarli
Department of Neurology, University of California, Davis, Sacramento, CA, USA.
Roles: Formal analysis, Funding acquisition, Investigation, Methodology, Resources, Software, Validation, and Writing - review & editing.
Myriam Fornage
Institute of Molecular Medicine, University of Texas Health Science Center at Houston, Houston, TX, USA.
Roles: Formal analysis, Resources, and Writing - review & editing.
Vilmundur Gudnason
Icelandic Heart Association, Kopavogur, Iceland.
Faculty of Medicine, University of Iceland, Reykjavik, Iceland.
Roles: Investigation, Resources, Supervision, and Writing - review & editing.
Edith Hofer
Clinical Division of Neurogeriatrics, Department of Neurology, Medical University of Graz, Graz, Austria.
Institute for Medical Informatics, Statistics and Documentation, Medical University of Graz, Graz, Austria.
Role: Validation.
M. Arfan Ikram
Department of Epidemiology, Erasmus MC Medical Center, Rotterdam, Netherlands.
Roles: Conceptualization, Data curation, Funding acquisition, Methodology, Project administration, and Resources.
Lenore Launer
Laboratory of Epidemiology and Population Sciences, Intramural Research Program, National Institute on Aging, Bethesda, MD, USA.
Roles: Methodology, Resources, and Writing - review & editing.
W. T. Longstreth
Departments of Neurology and Epidemiology, University of Washington, Seattle, WA, USA.
Roles: Resources and Writing - review & editing.
Department of Epidemiology, Erasmus MC Medical Center, Rotterdam, Netherlands.
Medical Informatics, Erasmus MC Medical Center, Rotterdam, Netherlands.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Visualization, and Writing - review & editing.
Neurodegeneratives Diseases Institute UMR 5293, CNRS, CEA, University of Bordeaux, Bordeaux, France.
Roles: Formal analysis, Investigation, Resources, and Software.
MIND Center, University of Mississippi Medical Center, Jackson, MS, USA.
Roles: Data curation, Funding acquisition, Investigation, Resources, and Writing - review & editing.
Gennady V. Roshchupkin
Department of Epidemiology, Erasmus MC Medical Center, Rotterdam, Netherlands.
Department of Radiology and Nuclear Medicine, Erasmus MC Medical Center, Rotterdam, Netherlands.
Medical Informatics, Erasmus MC Medical Center, Rotterdam, Netherlands.
Roles: Formal analysis, Investigation, Software, and Validation.
Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, University of Texas Health Sciences Center, San Antonio, TX, USA.
Department of Epidemiology & Biostatistics, University of Texas Health Sciences Center, San Antonio, TX, USA.
Department of Neurology, Boston University School of Medicine, Boston, MA, USA.
Roles: Data curation, Formal analysis, and Writing - review & editing.
Reinhold Schmidt
Clinical Division of Neurogeriatrics, Department of Neurology, Medical University of Graz, Graz, Austria.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Resources, Validation, and Writing - review & editing.
Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, University of Texas Health Sciences Center, San Antonio, TX, USA.
Department of Neurology, Boston University School of Medicine, Boston, MA, USA.
Framingham Heart Study and Department of Neurology, Boston University School of Medicine, Boston, MA, USA.
Roles: Conceptualization, Data curation, Funding acquisition, Project administration, Resources, Supervision, and Writing - review & editing.
Qiong Yang
Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA.
Roles: Formal analysis and Software.
Alzheimer’s Disease Neuroimaging Initiative
CHARGE Consortium
EPIGEN Consortium
IMAGEN Consortium
SYS Consortium
Parkinson’s Progression Markers Initiative
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Department of Neurology, FCM, UNICAMP, Campinas, Brazil.
Roles: Data curation, Formal analysis, Investigation, Resources, Validation, and Writing - review & editing.
David Ames
Academic Unit for Psychiatry of Old Age, University of Melbourne, Melbourne, VIC, Australia.
National Ageing Research Institute, Melbourne, VIC, Australia.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Visualization, and Writing - review & editing.
Department of Medicine, University of Otago, Christchurch, Christchurch, New Zealand.
New Zealand Brain Research Institute, Christchurch, New Zealand.
Brain Research New Zealand - Rangahau Roro Aotearoa, Christchurch, New Zealand.
Department of Neurology, Canterbury District Health Board, Christchurch, New Zealand.
Roles: Investigation and Resources.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Roles: Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, and Writing - review & editing.
Alejandro Arias-Vasquez
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Psychiatry, Radboud university medical center, Nijmegen, Netherlands.
Roles: Conceptualization, Methodology, Visualization, and Writing - review & editing.
Centre for Clinical Brain Sciences and Edinburgh Imaging, University of Edinburgh, Edinburgh, UK.
Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.
Roles: Data curation, Formal analysis, Investigation, and Resources.
Bernhard T. Baune
Department of Psychiatry, University of Münster, Münster, Germany.
Department of Psychiatry, Melbourne Medical School, University of Melbourne, Melbourne, VIC, Australia.
Florey Institute of Neuroscience and Mental Health, University of Melbourne, Parkville, VIC, Australia.
Roles: Conceptualization, Data curation, Investigation, Project administration, Resources, Supervision, Validation, Visualization, and Writing - review & editing.
Department of Psychiatry and Behavioral Sciences, Duke University School of Medicine, Durham, NC, USA.
VA Mid-Atlantic Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham, VA Healthcare System, Durham, NC, USA.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Project administration, Resources, and Supervision.
John Blangero
Department of Human Genetics and South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine, Brownsville, TX, USA.
Roles: Conceptualization, Formal analysis, Funding acquisition, Investigation, Methodology, Resources, Software, and Writing - review & editing.
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Funding acquisition, Investigation, Resources, Validation, and Writing - review & editing.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Dementia Centre for Research Collaboration, University of New South Wales, Sydney, NSW, Australia.
Roles: Funding acquisition, Investigation, and Writing - review & editing.
Han G. Brunner
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Clinical Genetics and School for Oncology and Developmental Biology (GROW), Maastricht University Medical Center, Maastricht, Netherlands.
Roles: Conceptualization, Investigation, Methodology, Project administration, Resources, Validation, and Writing - review & editing.
Randy L. Buckner
Department of Psychology and Center for Brain Science, Harvard University, Boston, MA, USA.
Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.
Department of Radiology, Massachusetts General Hospital, Boston, MA, USA.
Roles: Conceptualization, Data curation, Investigation, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.
Karakter Child and Adolescent Psychiatry University Center, Nijmegen, Netherlands.
Roles: Data curation, Methodology, Resources, Validation, Visualization, and Writing - review & editing.
Department of Psychiatry, University of New Mexico, Albuquerque, NM, USA.
Roles: Funding acquisition, Investigation, Resources, and Writing - review & editing.
Department of Psychiatry, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Funding acquisition, Investigation, Resources, and Validation.
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.
Schizophrenia Research Institute, Randwick, NSW, Australia.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, and Supervision.
Tri-institutional Center for Translational Research in Neuroimaging and Data Science (TReNDS), Georgia State University, Georgia Institute of Technology, Emory University, Atlanta, GA, USA.
Roles: Investigation, Resources, and Writing - review & editing.
School of Psychiatry, University of New South Wales, Sydney, NSW, Australia.
Neuroscience Research Australia, Sydney, NSW, Australia.
Department of Psychiatry, Monash University, Clayton, VIC, Australia.
Role: Resources.
Xavier Caseras
MRC Centre for Neuropsychiatric Genetics and Genomics, Cardiff University, Cardiff, UK.
Roles: Formal analysis, Investigation, and Writing - review & editing.
Svenja Caspers
Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.
JARA-BRAIN, Jülich-Aachen Research Alliance, Jülich, Germany.
Institute for Anatomy I, Medical Faculty, Heinrich-Heine University, Düsseldorf, Germany.
Roles: Investigation, Resources, and Writing - review & editing.
Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.
The SFI FutureNeuro Research Centre, Dublin, Ireland.
Roles: Funding acquisition, Investigation, Resources, Supervision, and Writing - review & editing.
Fernando Cendes
BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.
Department of Neurology, FCM, UNICAMP, Campinas, Brazil.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Supervision, and Validation.
Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Writing - original draft, and Writing - review & editing.
Benedicto Crespo-Facorro https://orcid.org/0000-0003-0033-7132
Centro Investigacion Biomedica en Red Salud Mental, Santander, Spain.
Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria–IDIVAL, Santander, Spain.
Hospital Universitario Virgen Del Rocio, IBiS, Universidad De Sevilla, Sevilla, Spain.
Roles: Conceptualization, Funding acquisition, Investigation, Resources, and Writing - review & editing.
John C. Dalrymple-Alford https://orcid.org/0000-0002-6672-033X
New Zealand Brain Research Institute, Christchurch, New Zealand.
Brain Research New Zealand - Rangahau Roro Aotearoa, Christchurch, New Zealand.
School of Psychology, Speech and Hearing, University of Canterbury, Christchurch, New Zealand.
Roles: Funding acquisition and Investigation.
Department of Psychiatry, University of Münster, Münster, Germany.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, and Validation.
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Formal analysis, Funding acquisition, Investigation, Project administration, Resources, and Writing - review & editing.
Ian J. Deary
Department of Psychology, University of Edinburgh, Edinburgh, UK.
Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.
Roles: Data curation, Funding acquisition, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Norman Delanty
Department of Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.
Future Neuro, Royal College of Surgeons in Ireland, Dublin, Ireland.
Roles: Conceptualization, Resources, and Validation.
Department of Neurology, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium.
Roles: Investigation, Resources, and Writing - review & editing.
Centre for Population Neuroscience and Precision Medicine (PONS), Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK.
Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK.
Roles: Conceptualization, Investigation, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Centre for Neuroimaging and Cognitive Genomics, School of Psychology, National University of Ireland Galway, Galway, Ireland.
Roles: Data curation, Investigation, Resources, Supervision, and Writing - review & editing.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.
Department of Psychology, University of Oslo, Oslo, Norway.
Roles: Formal analysis, Investigation, and Resources.
Guillén Fernández
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Project administration, Resources, and Supervision.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Roles: Conceptualization, Funding acquisition, Project administration, Resources, Supervision, and Writing - review & editing.
Department of Cognitive and Clinical Neuroscience, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.
Roles: Conceptualization, Funding acquisition, Methodology, Project administration, and Resources.
Andreas J. Forstner
Department of Biomedicine, University of Basel, Basel, Switzerland.
Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.
Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany.
Department of Psychiatry (UPK), University of Basel, Basel, Switzerland.
Centre for Human Genetics, University of Marburg, Marburg, Germany.
Roles: Formal analysis, Investigation, Resources, and Writing - review & editing.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
Roles: Resources and Writing - review & editing.
Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Department of Psychiatry, Radboud university medical center, Nijmegen, Netherlands.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Resources, Supervision, and Writing - review & editing.
David C. Glahn
Olin Neuropsychiatric Research Center, Institute of Living, Hartford Hospital, Hartford, CT, USA.
Tommy Fuss Center for Neuropsychiatric Disease Research, Boston Children’s Hospital and Department of Psychiatry, Harvard Medical School, Boston, MA, USA.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Supervision, and Writing - original draft.
Department of Psychiatry, Harvard Medical School, Boston, MA, USA.
Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.
Department of Radiology, Massachusetts General Hospital, Boston, MA, USA.
Roles: Conceptualization, Data curation, and Resources.
Hans J. Grabe
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany.
German Center for Neurodegenerative Diseases Rostock/Greifswald, Greifswald, Germany.
Roles: Conceptualization, Data curation, Investigation, Resources, Validation, and Writing - review & editing.
Oliver Gruber
Section for Experimental Psychopathology and Neuroimaging, Department of General Psychiatry, Heidelberg University Hospital, Heidelberg, Germany.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Supervision, and Validation.
Department of Neuroscience, Norwegian University of Science and Technology, Trondheim, Norway.
Department of Radiology and Nuclear Medicine, St. Olavs University Hospital, Trondheim, Norway.
Roles: Conceptualization, Data curation, Investigation, Project administration, Resources, Supervision, and Writing - review & editing.
Ahmad R. Hariri
Department of Psychology and Neuroscience, Duke University, Durham, NC, USA.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Resources, Supervision, and Writing - review & editing.
University of Groningen, University Medical Center Groningen, Department of Psychiatry, Groningen, Netherlands.
Roles: Investigation and Resources.
Ryota Hashimoto
Department of Psychiatry, Osaka University Graduate School of Medicine, Suita, Japan.
Molecular Research Center for Children’s Mental Development, United Graduate School of Child Development, Osaka University, Suita, Japan.
Department of Pathology of Mental Diseases, National Institute of Mental Health, National Center of Neurology and Psychiatry, Tokyo, Japan.
Roles: Investigation and Resources.
Andreas Heinz
Department of Psychiatry and Psychotherapy, Charité Campus Mitte, Charité - Universitätsmedizin Berlin, Berlin, Germany.
Roles: Conceptualization, Data curation, Methodology, Project administration, Supervision, Validation, Visualization, and Writing - review & editing.
School of Medicine and Public Health, University of Newcastle, Callaghan, NSW, Australia.
Health Behaviour Research Group, University of Newcastle, Callaghan, NSW, Australia.
Roles: Conceptualization, Funding acquisition, Investigation, Methodology, Supervision, and Writing - review & editing.
Department of Child and Adolescent Psychiatry/Psychology, Erasmus Medical Center-Sophia Children’s Hospital, Rotterdam, Netherlands.
Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Role: Resources.
Department of Psychiatry, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.
Roles: Resources and Writing - review & editing.
Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.
Department of Psychology, Yale University, New Haven, CT, USA.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Writing - original draft, and Writing - review & editing.
L. Elliot Hong
Maryland Psychiatry Research Center, Department of Psychiatry, University of Maryland School of Medicine, Baltimore, MD, USA.
Roles: Funding acquisition, Investigation, Project administration, and Resources.
William D. Hopkins
Department of Comparative Medicine, The University of Texas MD Anderson Cancer Center, Bastrop, TX, USA.
Role: Resources.
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Roles: Data curation, Investigation, and Resources.
Terry L. Jernigan
Department of Radiology, University of California San Diego, San Diego, CA, USA.
Department of Cognitive Science, University of California San Diego, San Diego, CA, USA.
Department of Psychiatry, University of California San Diego, San Diego, CA, USA.
Center for Human Development, University of California San Diego, La Jolla, CA, USA.
Roles: Conceptualization, Data curation, Investigation, and Project administration.
NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.
Centre for Psychiatric Research, Department of Clinical Neuroscience, Karolinska Institutet, Stockholm, Sweden.
Roles: Investigation and Resources.
René S. Kahn
Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
Roles: Conceptualization, Project administration, Resources, Supervision, and Writing - review & editing.
Department of Pathology and Biomedical Science, University of Otago, Christchurch, Christchurch, New Zealand.
Roles: Investigation, Resources, Supervision, and Writing - review & editing.
Department of Psychiatry and Psychotherapy, Philipps-University Marburg, Marburg, Germany.
Roles: Funding acquisition, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Peter Kochunov
Maryland Psychiatry Research Center, Department of Psychiatry, University of Maryland School of Medicine, Baltimore, MD, USA.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Software, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Neuroscience Research Australia, Sydney, NSW, Australia.
School of Medical Sciences, University of New South Wales, Sydney, NSW, Australia.
Neurogenetics and Epigenetics, Brain and Mind Centre, The University of Sydney, Sydney, NSW, Australia.
Roles: Investigation, Resources, and Supervision.
NORMENT, Department of Clinical Science, University of Bergen, Bergen, Norway.
Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway.
Roles: Formal analysis, Investigation, Resources, and Validation.
Carmel M. Loughland
School of Medicine and Public Health, University of Newcastle, Callaghan, NSW, Australia.
Hunter New England Mental Health Service, Newcastle, NSW, Australia.
Roles: Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, Visualization, and Writing - review & editing.
Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
Roles: Conceptualization, Data curation, and Resources.
Jean-Luc Martinot
INSERM Unit 1000 - Neuroimaging & Psychiatry, Paris Saclay University, Gif sur Yvette, France.
Roles: Funding acquisition, Investigation, Resources, and Supervision.
Colm McDonald
Centre for Neuroimaging and Cognitive Genomics, National University of Ireland Galway, Galway, Ireland.
Roles: Funding acquisition, Investigation, and Writing - review & editing.
Herston Imaging Research Facility, School of Clinical Sciences, Queensland University of Technology, Brisbane, QLD, Australia.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Resources, and Writing - review & editing.
Andreas Meyer-Lindenberg https://orcid.org/0000-0001-5619-1123
Department of Psychiatry and Psychotherapy, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.
Roles: Conceptualization and Writing - review & editing.
School of Psychology, University of Newcastle, Callaghan, NSW, Australia.
Roles: Funding acquisition, Investigation, Project administration, and Supervision.
Duke UNC Brain Imaging and Analysis Center, Duke University Medical Center, Durham, NC, USA.
Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham VA Medical Center, Durham, NC, USA.
Roles: Investigation, Resources, and Writing - review & editing.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Queensland Centre for Mental Health Research, University of Queensland, Brisbane, QLD, Australia.
Roles: Conceptualization, Funding acquisition, Investigation, Project administration, Resources, and Writing - review & editing.
Lars Nyberg
Department of Integrative Medical Biology, Umeå University, Umeå, Sweden.
Umeå Center for Functional Brain Imaging, Umeå University, Umeå, Sweden.
Department of Radiation Sciences, Umeå University, Umeå, Sweden.
Roles: Data curation, Formal analysis, Project administration, and Writing - review & editing.
Emma Children's Hospital Academic Medical Center, Amsterdam, Netherlands.
Department of Pediatrics, Vrije Universiteit Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Clinical Neuropsychology section, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Data curation, Investigation, Project administration, Resources, and Writing - review & editing.
Center for Neurobehavioral Genetics, University of California Los Angeles, Los Angeles, CA, USA.
Roles: Data curation, Investigation, Resources, Validation, and Writing - review & editing.
Department of Psychiatry, Melbourne Medical School, University of Melbourne, Melbourne, VIC, Australia.
Florey Institute of Neuroscience and Mental Health, University of Melbourne, Parkville, VIC, Australia.
NorthWestern Mental Health, Sunshine Hospital, St Albans, VIC, Australia.
Roles: Data curation, Funding acquisition, Investigation, Resources, Supervision, Validation, and Writing - review & editing.
Tomas Paus
Bloorview Research Institute, University of Toronto, Toronto, ON, Canada.
Departments of Psychology and Psychiatry, University of Toronto, Toronto, ON, Canada.
Centre for Developing Brain, Child Mind Institute, New York, NY, USA.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Writing - original draft, and Writing - review & editing.
The Hospital for Sick Children, University of Toronto, Toronto, ON, Canada.
Department of Physiology, University of Toronto, Toronto, ON, Canada.
Roles: Conceptualization, Funding acquisition, Investigation, Project administration, Supervision, Validation, and Writing - review & editing.
Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, Netherlands.
Roles: Conceptualization, Data curation, Investigation, Resources, Validation, and Writing - review & editing.
Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Formal analysis, Investigation, and Resources.
Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Department of Clinical Genetics, Vrije Universiteit Medical Centre, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Formal analysis, Investigation, Resources, and Writing - review & editing.
Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.
Roles: Conceptualization, Funding acquisition, Resources, Validation, and Writing - review & editing.
Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.
Roles: Conceptualization, Investigation, Resources, and Writing - review & editing.
Laura M. Rowland
Maryland Psychiatry Research Center, Department of Psychiatry, University of Maryland School of Medicine, Baltimore, MD, USA.
Roles: Investigation and Writing - review & editing.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Neuropsychiatric Institute, The Prince of Wales Hospital, Sydney, NSW, Australia.
Roles: Funding acquisition, Project administration, Resources, Supervision, and Writing - review & editing.
Philipp G. Sämann
Max Planck Institute of Psychiatry, Munich, Germany.
Roles: Formal analysis, Investigation, and Resources.
Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.
Hunter Medical Research Institute, Newcastle, NSW, Australia.
Roles: Funding acquisition, Investigation, and Resources.
Gunter Schumann
Institute of Science and Technology for Brain-Inspired Intelligence, Fudan University, Shanghai, China.
Centre for Population Neuroscience and Precision Medicine (PONS), Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK.
Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK.
PONS Research Group, Department of Psychiatry and Psychotherapie, Charité Campus Mitte, Humboldt University Berlin, Berlin, Germany.
Leibniz Institute for Neurobiology, Magdeburg, Germany.
Roles: Conceptualization, Formal analysis, Funding acquisition, Project administration, Resources, and Supervision.
School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.
Division of Molecular Medicine, John Hunter Hospital, New Lambton Heights, NSW, Australia.
Roles: Investigation, Resources, and Validation.
General Psychiatry, Institute of Mental Health, Singapore, Singapore.
Roles: Data curation, Formal analysis, Funding acquisition, Resources, and Writing - review & editing.
Sanjay M. Sisodiya
Department of Clinical and Experimental Epilepsy, UCL Queen Square Institute of Neurology, London, UK.
Chalfont Centre for Epilepsy, Chalfont-St-Peter, UK.
Roles: Investigation, Supervision, and Writing - review & editing.
Jordan W. Smoller
Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA.
Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.
Stanley Center for Psychiatric Research, Broad Institute, Boston, MA, USA.
Roles: Project administration and Supervision.
Cognitive Neuroscience Center, Department of Biomedical Sciences of Cells and Systems, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.
Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.
Department of Psychiatry, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.
Department of Medical and Biological Psychology, University of Bergen, Bergen, Norway.
Roles: Conceptualization, Data curation, Formal analysis, Investigation, Resources, Writing - original draft, and Writing - review & editing.
Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.
Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.
MRC Integrative Epidemiology Unit, Department of Population Health Sciences, Bristol Medical School, Bristol, UK.
Roles: Formal analysis, Investigation, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Department of Psychiatry and Neuroscience Institute, University of Cape Town, Cape Town, South Africa.
MRC Unit on Risk & Resilience in Mental Disorders, University of Cape Town, Cape Town, South Africa.
Roles: Conceptualization, Funding acquisition, Investigation, and Writing - review & editing.
Laboratory of Neuro Imaging, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of the University of Southern California, Los Angeles, CA, USA.
Roles: Resources, Supervision, and Writing - review & editing.
Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.
Department of Developmental Disability Neuropsychiatry, University of New South Wales, Sydney, NSW, Australia.
Roles: Conceptualization, Data curation, Funding acquisition, Investigation, Project administration, Supervision, Validation, and Writing - review & editing.
Department of Psychiatry, Leiden University Medical Center, Leiden, Netherlands.
Roles: Investigation, Resources, and Writing - review & editing.
Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
Roles: Data curation, Formal analysis, Funding acquisition, Investigation, Resources, Software, Validation, and Writing - review & editing.
Henry Völzke
Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.
Roles: Conceptualization, Funding acquisition, Investigation, and Resources.
Division of Mind and Brain Research, Department of Psychiatry and Psychotherapy, Campus Charité Mitte, Charité - Universitätsmedizin Berlin corporate member of Freie Universität Berlin, Humboldt-Universität zu Berlin, and Berlin Institute of Health, Berlin, Germany.
Roles: Conceptualization, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, and Writing - review & editing.
Center for Economics and Neuroscience, University of Bonn, Bonn, Germany.
Institute of Experimental Epileptology and Cognition Research, University Hospital Bonn, Germany.
Roles: Data curation, Investigation, and Resources.
Lieber Institute for Brain Development, Baltimore, MD, USA.
Psychiatry, Neurology, Neuroscience, Genetics, Johns Hopkins University, Baltimore, MD, USA.
Role: Validation.
Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.
Centre for Advanced Imaging, University of Queensland, Brisbane, QLD, Australia.
Roles: Conceptualization, Data curation, Investigation, Resources, Supervision, and Writing - review & editing.
Center for Cognitive Neuroscience, Neuroscience and Behavioral Disorders Program, Duke–National University of Singapore Medical School, Singapore, Singapore.
Roles: Formal analysis, Investigation, Software, and Writing - review & editing.
Department of Genetics and UNC Neuroscience Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Imaging Genetics Center, Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of USC, University of Southern California, Los Angeles, CA, USA.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.
School of Psychology, University of Queensland, Brisbane, QLD, Australia.
Faculty of Medicine, University of Queensland, Brisbane, QLD, Australia.
Roles: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, and Writing - review & editing.
Enhancing NeuroImaging Genetics through Meta-Analysis Consortium (ENIGMA)—Genetics working group

Correction

21 October 2021: See Erratum at science.org/doi/10.1126/science.abm7211. The original version is available here:

Funding Information

NIH Clinical Center: R01MH084898-01A1
UK Medical Research Council
South-Eastern Norway Regional Health Authority: 2017112
The Research Council of Norway: 223273
KG Jebsen Foundation: SKGJ‐MED‐008
FAPESP - São Paulo Research Foundation - Brazil: 2013/07559-3
Tond Mohn Stiftelse: Career grant to SLH

Notes

*
Corresponding author. Email: [email protected] (K.L.G.); [email protected] (N.J.); [email protected] (J.L.S.); [email protected] (P.M.T.); [email protected] (S.E.M.)
These authors contributed equally to this work.
These authors contributed equally to this work.
§
These authors contributed equally to this work.
Consortium authors are listed in the supplementary materials.

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