The genetic architecture of the human cerebral cortex-Reference-Cited by-同舟云学术

The genetic architecture of the human cerebral cortex

Published:2020-03-20 Issue:6484 Volume:367 Page:
ISSN:0036-8075
Container-title:Science
language:en
Short-container-title:Science

Author:

Grasby Katrina L.¹^ORCID,Jahanshad Neda²^ORCID,Painter Jodie N.¹^ORCID,Colodro-Conde Lucía¹³⁴⁵^ORCID,Bralten Janita⁶⁷^ORCID,Hibar Derrek P.²⁸,Lind Penelope A.¹⁴⁹^ORCID,Pizzagalli Fabrizio²,Ching Christopher R. K.²¹⁰,McMahon Mary Agnes B.²^ORCID,Shatokhina Natalia²,Zsembik Leo C. P.¹¹^ORCID,Thomopoulos Sophia I.²,Zhu Alyssa H.²^ORCID,Strike Lachlan T.¹²^ORCID,Agartz Ingrid¹³¹⁴¹⁵¹⁶^ORCID,Alhusaini Saud¹⁷¹⁸,Almeida Marcio A. A.¹⁹^ORCID,Alnæs Dag¹³¹⁴,Amlien Inge K.²⁰^ORCID,Andersson Micael²¹²²^ORCID,Ard Tyler²³^ORCID,Armstrong Nicola J.²⁴^ORCID,Ashley-Koch Allison²⁵,Atkins Joshua R.²⁶²⁷^ORCID,Bernard Manon²⁸^ORCID,Brouwer Rachel M.²⁹^ORCID,Buimer Elizabeth E. L.²⁹^ORCID,Bülow Robin³⁰,Bürger Christian³¹,Cannon Dara M.³²,Chakravarty Mallar³³³⁴,Chen Qiang³⁵^ORCID,Cheung Joshua W.²^ORCID,Couvy-Duchesne Baptiste¹²³⁶³⁷^ORCID,Dale Anders M.³⁸³⁹^ORCID,Dalvie Shareefa⁴⁰,de Araujo Tânia K.⁴¹⁴²^ORCID,de Zubicaray Greig I.⁴³^ORCID,de Zwarte Sonja M. C.²⁹,den Braber Anouk⁴⁴⁴⁵,Doan Nhat Trung¹³¹⁴,Dohm Katharina³¹^ORCID,Ehrlich Stefan⁴⁶^ORCID,Engelbrecht Hannah-Ruth⁴⁷^ORCID,Erk Susanne⁴⁸,Fan Chun Chieh⁴⁹,Fedko Iryna O.⁴⁴^ORCID,Foley Sonya F.⁵⁰,Ford Judith M.⁵¹^ORCID,Fukunaga Masaki⁵²^ORCID,Garrett Melanie E.²⁵^ORCID,Ge Tian⁵³⁵⁴,Giddaluru Sudheer⁵⁵,Goldman Aaron L.³⁵,Green Melissa J.⁵⁶⁵⁷^ORCID,Groenewold Nynke A.⁴⁰,Grotegerd Dominik³¹^ORCID,Gurholt Tiril P.¹³¹⁴¹⁵^ORCID,Gutman Boris A.²⁵⁸^ORCID,Hansell Narelle K.¹²^ORCID,Harris Mathew A.⁵⁹⁶⁰^ORCID,Harrison Marc B.²^ORCID,Haswell Courtney C.⁶¹⁶²^ORCID,Hauser Michael²⁵^ORCID,Herms Stefan⁶³⁶⁴⁶⁵^ORCID,Heslenfeld Dirk J.⁶⁶,Ho New Fei⁶⁷,Hoehn David⁶⁸^ORCID,Hoffmann Per⁶³⁶⁴⁶⁹^ORCID,Holleran Laurena⁷⁰^ORCID,Hoogman Martine⁶⁷^ORCID,Hottenga Jouke-Jan⁴⁴^ORCID,Ikeda Masashi⁷¹^ORCID,Janowitz Deborah⁷²^ORCID,Jansen Iris E.⁷³⁷⁴^ORCID,Jia Tianye⁷⁵⁷⁶⁷⁷^ORCID,Jockwitz Christiane⁷⁸⁷⁹⁸⁰^ORCID,Kanai Ryota⁸¹⁸²⁸³^ORCID,Karama Sherif³³⁸⁴⁸⁵^ORCID,Kasperaviciute Dalia⁸⁶⁸⁷,Kaufmann Tobias¹³¹⁴^ORCID,Kelly Sinead⁸⁸⁸⁹,Kikuchi Masataka⁹⁰^ORCID,Klein Marieke⁶⁷²⁹^ORCID,Knapp Michael⁹¹,Knodt Annchen R.⁹²^ORCID,Krämer Bernd⁹³⁹⁴^ORCID,Lam Max⁶⁷⁹⁵,Lancaster Thomas M.⁵⁰⁹⁶,Lee Phil H.⁵³⁹⁷^ORCID,Lett Tristram A.⁴⁸,Lewis Lindsay B.⁸⁵⁹⁸^ORCID,Lopes-Cendes Iscia⁴¹⁴²^ORCID,Luciano Michelle⁹⁹¹⁰⁰^ORCID,Macciardi Fabio¹⁰¹^ORCID,Marquand Andre F.⁷¹⁰²^ORCID,Mathias Samuel R.¹⁰³¹⁰⁴,Melzer Tracy R.¹⁰⁵¹⁰⁶¹⁰⁷^ORCID,Milaneschi Yuri¹⁰⁸,Mirza-Schreiber Nazanin⁶⁸¹⁰⁹^ORCID,Moreira Jose C. V.⁴²¹¹⁰^ORCID,Mühleisen Thomas W.⁶³⁷⁸¹¹¹^ORCID,Müller-Myhsok Bertram⁶⁸¹¹²¹¹³,Najt Pablo³²^ORCID,Nakahara Soichiro¹⁰¹¹¹⁴,Nho Kwangsik¹¹⁵,Olde Loohuis Loes M.¹¹⁶^ORCID,Orfanos Dimitri Papadopoulos¹¹⁷^ORCID,Pearson John F.¹¹⁸¹¹⁹^ORCID,Pitcher Toni L.¹⁰⁵¹⁰⁶¹⁰⁷^ORCID,Pütz Benno⁶⁸,Quidé Yann⁵⁶⁵⁷^ORCID,Ragothaman Anjanibhargavi²^ORCID,Rashid Faisal M.²,Reay William R.²⁶²⁷^ORCID,Redlich Ronny³¹,Reinbold Céline S.²⁰⁶³⁶⁴^ORCID,Repple Jonathan³¹^ORCID,Richard Geneviève¹³¹⁴¹²⁰¹²¹^ORCID,Riedel Brandalyn C.²¹¹⁵^ORCID,Risacher Shannon L.¹¹⁵,Rocha Cristiane S.⁴¹⁴²^ORCID,Mota Nina Roth⁶⁷¹²²^ORCID,Salminen Lauren²^ORCID,Saremi Arvin²^ORCID,Saykin Andrew J.¹¹⁵¹²³^ORCID,Schlag Fenja¹²⁴^ORCID,Schmaal Lianne¹²⁵¹²⁶¹²⁷,Schofield Peter R.⁵⁷¹²⁸^ORCID,Secolin Rodrigo⁴¹⁴²^ORCID,Shapland Chin Yang¹²⁴^ORCID,Shen Li¹²⁹^ORCID,Shin Jean²⁸¹³⁰^ORCID,Shumskaya Elena⁶⁷¹³¹^ORCID,Sønderby Ida E.¹³¹⁴,Sprooten Emma⁷^ORCID,Tansey Katherine E.⁹⁶,Teumer Alexander¹³²,Thalamuthu Anbupalam¹³³^ORCID,Tordesillas-Gutiérrez Diana¹³⁴¹³⁵,Turner Jessica A.¹³⁶¹³⁷^ORCID,Uhlmann Anne⁴⁰¹³⁸,Vallerga Costanza Ludovica³⁶^ORCID,van der Meer Dennis¹³⁹¹⁴⁰,van Donkelaar Marjolein M. J.¹⁴¹,van Eijk Liza³¹²^ORCID,van Erp Theo G. M.¹⁰¹^ORCID,van Haren Neeltje E. M.²⁹¹⁴²^ORCID,van Rooij Daan⁷¹⁰²^ORCID,van Tol Marie-José¹⁴³^ORCID,Veldink Jan H.¹⁴⁴^ORCID,Verhoef Ellen¹²⁴^ORCID,Walton Esther¹³⁶¹⁴⁵¹⁴⁶^ORCID,Wang Mingyuan⁶⁷^ORCID,Wang Yunpeng¹³¹⁴^ORCID,Wardlaw Joanna M.⁵⁹¹⁰⁰¹⁴⁷^ORCID,Wen Wei¹³³^ORCID,Westlye Lars T.¹³¹⁴¹²⁰^ORCID,Whelan Christopher D.²¹⁷,Witt Stephanie H.¹⁴⁸^ORCID,Wittfeld Katharina⁷²¹⁴⁹^ORCID,Wolf Christiane¹⁵⁰^ORCID,Wolfers Thomas⁶^ORCID,Wu Jing Qin²⁶^ORCID,Yasuda Clarissa L.⁴²¹⁵¹^ORCID,Zaremba Dario³¹,Zhang Zuo¹⁵²^ORCID,Zwiers Marcel P.⁷¹⁰²¹³¹,Artiges Eric¹⁵³^ORCID,Assareh Amelia A.¹³³,Ayesa-Arriola Rosa¹³⁵¹⁵⁴^ORCID,Belger Aysenil⁶¹¹⁵⁵^ORCID,Brandt Christine L.¹³¹⁴,Brown Gregory G.¹⁵⁶¹⁵⁷,Cichon Sven⁶³⁶⁴⁷⁸^ORCID,Curran Joanne E.¹⁹^ORCID,Davies Gareth E.¹⁵⁸,Degenhardt Franziska⁶⁹,Dennis Michelle F.⁶²^ORCID,Dietsche Bruno¹⁵⁹,Djurovic Srdjan¹⁶⁰¹⁶¹^ORCID,Doherty Colin P.¹⁶²¹⁶³¹⁶⁴^ORCID,Espiritu Ryan¹⁶⁵,Garijo Daniel¹⁶⁵^ORCID,Gil Yolanda¹⁶⁵,Gowland Penny A.¹⁶⁶^ORCID,Green Robert C.¹⁶⁷¹⁶⁸¹⁶⁹^ORCID,Häusler Alexander N.¹⁷⁰¹⁷¹^ORCID,Heindel Walter¹⁷²,Ho Beng-Choon¹⁷³^ORCID,Hoffmann Wolfgang U.¹³²¹⁴⁹^ORCID,Holsboer Florian⁶⁸¹⁷⁴^ORCID,Homuth Georg¹⁷⁵^ORCID,Hosten Norbert¹⁷⁶^ORCID,Jack Clifford R.¹⁷⁷^ORCID,Jang MiHyun¹⁶⁵^ORCID,Jansen Andreas¹⁵⁹¹⁷⁸^ORCID,Kimbrel Nathan A.⁶²¹⁷⁹,Kolskår Knut¹³¹⁴¹²⁰¹²¹,Koops Sanne²⁹,Krug Axel¹⁵⁹^ORCID,Lim Kelvin O.¹⁸⁰^ORCID,Luykx Jurjen J.²⁹¹⁸¹¹⁸²^ORCID,Mathalon Daniel H.¹⁸³¹⁸⁴^ORCID,Mather Karen A.⁵⁷¹³³,Mattay Venkata S.³⁵¹⁸⁵¹⁸⁶^ORCID,Matthews Sarah¹⁴⁵,Mayoral Van Son Jaqueline¹³⁵¹⁵⁴,McEwen Sarah C.¹⁸⁷¹⁸⁸^ORCID,Melle Ingrid¹³¹⁴^ORCID,Morris Derek W.³²^ORCID,Mueller Bryon A.¹⁸⁰^ORCID,Nauck Matthias¹⁸⁹¹⁹⁰,Nordvik Jan E.¹²¹^ORCID,Nöthen Markus M.⁶⁹^ORCID,O’Leary Daniel S.¹⁷³^ORCID,Opel Nils³¹,Martinot Marie-Laure Paillère¹⁵³¹⁹¹,Pike G. Bruce¹⁹²,Preda Adrian¹⁹³^ORCID,Quinlan Erin B.¹⁵²,Rasser Paul E.²⁷¹⁹⁴¹⁹⁵¹⁹⁶^ORCID,Ratnakar Varun¹⁶⁵^ORCID,Reppermund Simone¹³³¹⁹⁷,Steen Vidar M.¹⁶¹¹⁹⁸^ORCID,Tooney Paul A.²⁶¹⁹⁶,Torres Fábio R.⁴¹⁴²^ORCID,Veltman Dick J.¹⁰⁸^ORCID,Voyvodic James T.⁶¹^ORCID,Whelan Robert¹⁹⁹^ORCID,White Tonya¹⁴²²⁰⁰,Yamamori Hidenaga²⁰¹^ORCID,Adams Hieab H. H.²⁰²²⁰³²⁰⁴^ORCID,Bis Joshua C.²⁰⁵,Debette Stephanie²⁰⁶²⁰⁷,Decarli Charles²⁰⁸,Fornage Myriam²⁰⁹,Gudnason Vilmundur²¹⁰²¹¹,Hofer Edith²¹²²¹³,Ikram M. Arfan²⁰²,Launer Lenore²¹⁴,Longstreth W. T.²¹⁵,Lopez Oscar L.²⁰²²¹⁶^ORCID,Mazoyer Bernard²¹⁷^ORCID,Mosley Thomas H.²¹⁸^ORCID,Roshchupkin Gennady V.²⁰²²⁰³²¹⁶,Satizabal Claudia L.²¹⁹²²⁰²²¹^ORCID,Schmidt Reinhold²¹²,Seshadri Sudha²¹⁹²²¹²²²^ORCID,Yang Qiong²²³,Alvim Marina K. M.⁴²¹⁵¹^ORCID,Ames David²²⁴²²⁵,Anderson Tim J.¹⁰⁵¹⁰⁶¹⁰⁷²²⁶^ORCID,Andreassen Ole A.¹³¹⁴^ORCID,Arias-Vasquez Alejandro⁶⁷¹²²,Bastin Mark E.⁵⁹¹⁰⁰^ORCID,Baune Bernhard T.³¹²²⁷²²⁸,Beckham Jean C.¹⁷⁹²²⁹^ORCID,Blangero John¹⁹,Boomsma Dorret I.⁴⁴^ORCID,Brodaty Henry¹³³²³⁰^ORCID,Brunner Han G.⁶⁷²³¹,Buckner Randy L.²³²²³³²³⁴,Buitelaar Jan K.⁷¹⁰²²³⁵^ORCID,Bustillo Juan R.²³⁶^ORCID,Cahn Wiepke²³⁷^ORCID,Cairns Murray J.²⁶²⁷²³⁸^ORCID,Calhoun Vince²³⁹^ORCID,Carr Vaughan J.⁵⁶⁵⁷²⁴⁰^ORCID,Caseras Xavier⁹⁶,Caspers Svenja⁷⁸⁸⁰²⁴¹,Cavalleri Gianpiero L.²⁴²²⁴³^ORCID,Cendes Fernando⁴²¹⁵¹,Corvin Aiden²⁴⁴^ORCID,Crespo-Facorro Benedicto¹³⁵¹⁵⁴²⁴⁵^ORCID,Dalrymple-Alford John C.¹⁰⁶¹⁰⁷²⁴⁶^ORCID,Dannlowski Udo³¹^ORCID,de Geus Eco J. C.⁴⁴^ORCID,Deary Ian J.⁹⁹¹⁰⁰,Delanty Norman¹⁷¹⁶⁴,Depondt Chantal²⁴⁷^ORCID,Desrivières Sylvane⁷⁷¹⁵²^ORCID,Donohoe Gary⁷⁰^ORCID,Espeseth Thomas¹³¹²⁰^ORCID,Fernández Guillén⁷¹⁰²,Fisher Simon E.⁷¹²⁴^ORCID,Flor Herta²⁴⁸^ORCID,Forstner Andreas J.⁶³⁶⁴⁶⁹²⁴⁹²⁵⁰,Francks Clyde⁷¹²⁴^ORCID,Franke Barbara⁶⁷¹²²^ORCID,Glahn David C.¹⁰⁴²⁵¹,Gollub Randy L.⁹⁷²³³²³⁴^ORCID,Grabe Hans J.⁷²¹⁴⁹,Gruber Oliver⁹³,Håberg Asta K.²⁵²²⁵³^ORCID,Hariri Ahmad R.⁹²,Hartman Catharina A.²⁵⁴^ORCID,Hashimoto Ryota²⁰¹²⁵⁵²⁵⁶,Heinz Andreas²⁵⁷,Henskens Frans A.¹⁹⁴²⁵⁸^ORCID,Hillegers Manon H. J.¹⁴²²⁵⁹^ORCID,Hoekstra Pieter J.²⁶⁰^ORCID,Holmes Avram J.²³³²⁶¹^ORCID,Hong L. Elliot²⁶²,Hopkins William D.²⁶³,Hulshoff Pol Hilleke E.²⁹^ORCID,Jernigan Terry L.³⁹⁴⁹¹⁵⁶²⁶⁴,Jönsson Erik G.¹⁴¹⁶^ORCID,Kahn René S.²⁹²⁶⁵,Kennedy Martin A.¹¹⁹^ORCID,Kircher Tilo T. J.¹⁵⁹^ORCID,Kochunov Peter²⁶²,Kwok John B. J.⁵⁷¹²⁸²⁶⁶^ORCID,Le Hellard Stephanie¹⁶¹¹⁹⁸^ORCID,Loughland Carmel M.¹⁹⁴²⁶⁷,Martin Nicholas G.³⁷^ORCID,Martinot Jean-Luc¹⁵³,McDonald Colm³²,McMahon Katie L.²⁶⁸^ORCID,Meyer-Lindenberg Andreas²⁶⁹^ORCID,Michie Patricia T.²⁷⁰^ORCID,Morey Rajendra A.⁶¹⁶²^ORCID,Mowry Bryan¹²²⁷¹^ORCID,Nyberg Lars²¹²²²⁷²,Oosterlaan Jaap²⁷³²⁷⁴²⁷⁵^ORCID,Ophoff Roel A.¹¹⁶^ORCID,Pantelis Christos²²⁷²²⁸²⁷⁶^ORCID,Paus Tomas²⁷⁷²⁷⁸²⁷⁹,Pausova Zdenka²⁸²⁸⁰^ORCID,Penninx Brenda W. J. H.¹⁰⁸^ORCID,Polderman Tinca J. C.⁷³^ORCID,Posthuma Danielle⁷³²⁸¹^ORCID,Rietschel Marcella¹⁴⁸^ORCID,Roffman Joshua L.²³³^ORCID,Rowland Laura M.²⁶²,Sachdev Perminder S.¹³³²⁸²^ORCID,Sämann Philipp G.⁶⁸,Schall Ulrich²⁷¹⁹⁶^ORCID,Schumann Gunter⁷⁵⁷⁷¹⁵²²⁸³²⁸⁴,Scott Rodney J.²⁶²⁸⁵^ORCID,Sim Kang²⁸⁶^ORCID,Sisodiya Sanjay M.⁸⁶²⁸⁷,Smoller Jordan W.⁵³²³³²⁸⁸,Sommer Iris E.¹⁴³²⁵⁹²⁶⁰²⁸⁹^ORCID,St Pourcain Beate⁷¹²⁴¹⁴⁵^ORCID,Stein Dan J.²⁹⁰²⁹¹^ORCID,Toga Arthur W.²³^ORCID,Trollor Julian N.¹³³¹⁹⁷^ORCID,Van der Wee Nic J. A.²⁹²^ORCID,van ’t Ent Dennis⁴⁴^ORCID,Völzke Henry¹³²,Walter Henrik⁴⁸^ORCID,Weber Bernd¹⁷⁰¹⁷¹^ORCID,Weinberger Daniel R.³⁵²⁹³^ORCID,Wright Margaret J.¹²²⁹⁴^ORCID,Zhou Juan²⁹⁵^ORCID,Stein Jason L.¹¹^ORCID,Thompson Paul M.²^ORCID,Medland Sarah E.¹³⁹^ORCID, , , , , , ,

Affiliation:

1. Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.

2. 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.

3. School of Psychology, University of Queensland, Brisbane, QLD, Australia.

4. School of Biomedical Sciences, Queensland University of Technology, Brisbane, QLD, Australia.

5. Faculty of Psychology, University of Murcia, Murcia, Spain.

6. Department of Human Genetics, Radboud University Medical Center, Nijmegen, Netherlands.

7. Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands.

8. Personalized Healthcare, Genentech, Inc., South San Francisco, CA, USA.

9. Faculty of Medicine, University of Queensland, Brisbane, QLD, Australia.

10. Graduate Interdepartmental Program in Neuroscience, University of California Los Angeles, Los Angeles, CA, USA.

11. Department of Genetics and UNC Neuroscience Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.

12. Queensland Brain Institute, University of Queensland, St Lucia, QLD, Australia.

13. NORMENT - K.G. Jebsen Centre for Psychosis Research, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.

14. NORMENT - K.G. Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

15. Department of Psychiatric Research, Diakonhjemmet Hospital, Oslo, Norway.

16. Centre for Psychiatric Research, Department of Clinical Neuroscience, Karolinska Institutet, Stockholm, Sweden.

17. Department of Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.

18. Neurology Department, Yale School of Medicine, New Haven, CT, USA.

19. Department of Human Genetics and South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine, Brownsville, TX, USA.

20. Centre for Lifespan Changes in Brain and Cognition, Department of Psychology, University of Oslo, Oslo, Norway.

21. Department of Integrative Medical Biology, Umeå University, Umeå, Sweden.

22. Umeå Center for Functional Brain Imaging, Umeå University, Umeå, Sweden.

23. 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.

24. Mathematics and Statistics, Murdoch University, Murdoch, WA, Australia.

25. Duke Molecular Physiology Institute, Duke University Medical Center, Durham, NC, USA.

26. School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, NSW, Australia.

27. Priority Centre for Brain and Mental Health Research, University of Newcastle, Callaghan, NSW, Australia.

28. The Hospital for Sick Children, University of Toronto, Toronto, ON, Canada.

29. Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.

30. Institute for Radiology and Neuroradiology, University Medicine, Ernst-Moritz-Arndt University, Greifswald, Germany.

31. Department of Psychiatry, University of Münster, Münster, Germany.

32. Centre for Neuroimaging and Cognitive Genomics, National University of Ireland Galway, Galway, Ireland.

33. Douglas Mental Health University Institute, McGill University, Montreal, QC, Canada.

34. Departments of Psychiatry and Biological and Biomedical Engineering, McGill University, Montreal, QC, Canada.

35. Lieber Institute for Brain Development, Baltimore, MD, USA.

36. Institute for Molecular Bioscience, University of Queensland, Brisbane, QLD, Australia.

37. Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia.

38. Department of Neurosciences, University of California, San Diego, La Jolla, CA, USA.

39. Department of Radiology, University of California San Diego, San Diego, CA, USA.

40. Department of Psychiatry and Mental Health, University of Cape Town, Cape Town, South Africa.

41. Department of Medical Genetics and Genomic Medicine, School of Medical Sciences, University of Campinas - UNICAMP, Campinas, Brazil.

42. BRAINN - Brazilian Institute of Neuroscience and Neurotechnology, Campinas, Brazil.

43. Faculty of Health, Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, QLD, Australia.

44. Department of Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

45. Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Vrije Universiteit Amsterdam, Amsterdam UMC, Amsterdam, Netherlands.

46. Division of Psychological and Social Medicine and Developmental Neurosciences, Faculty of Medicine, Technische Universität Dresden, Dresden, Germany.

47. Division of Human Genetics, Institute of Infectious Disease and Molecular Medicine, University of Cape Town, Cape Town, South Africa.

48. 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.

49. Department of Cognitive Science, University of California San Diego, San Diego, CA, USA.

50. Cardiff University Brain Research Imaging Centre, Cardiff University, Cardiff, UK.

51. San Francisco Veterans Administration Medical Center, San Francisco, CA, USA.

52. Division of Cerebral Integration, National Institute for Physiological Sciences, Okazaki, Japan.

53. Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA.

54. Athinoula A. Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Boston, MA, USA.

55. NORMENT - K.G. Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway.

56. School of Psychiatry, University of New South Wales, Sydney, NSW, Australia.

57. Neuroscience Research Australia, Sydney, NSW, Australia.

58. Department of Biomedical Engineering, Illinois Institute of Technology, Chicago, IL, USA.

59. Centre for Clinical Brain Sciences and Edinburgh Imaging, University of Edinburgh, Edinburgh, UK.

60. Division of Psychiatry, University of Edinburgh, Edinburgh, UK.

61. Duke UNC Brain Imaging and Analysis Center, Duke University Medical Center, Durham, NC, USA.

62. Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham VA Medical Center, Durham, NC, USA.

63. Department of Biomedicine, University of Basel, Basel, Switzerland.

64. Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland.

65. Department of Genomics, Life & Brain Research Center, University of Bonn, Bonn, Germany.

66. Department of Cognitive and Clinical Neuropsychology, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

67. Research Division, Institute of Mental Health, Singapore, Singapore.

68. Max Planck Institute of Psychiatry, Munich, Germany.

69. Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany.

70. Centre for Neuroimaging and Cognitive Genomics, School of Psychology, National University of Ireland Galway, Galway, Ireland.

71. Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Japan.

72. Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany.

73. Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

74. Department of Neurology, Alzheimer Center, Amsterdam Neuroscience, Vrije Universiteit Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

75. Institute of Science and Technology for Brain-Inspired Intelligence, Fudan University, Shanghai, China.

76. Key Laboratory of Computational Neuroscience and Brain-Inspired Intelligence (Fudan University), Ministry of Education, Shanghai, China.

77. Centre for Population Neuroscience and Precision Medicine (PONS), Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK.

78. Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.

79. Department of Psychiatry, Psychotherapy and Psychosomatics, Medical Faculty, RWTH Aachen University, Aachen, Germany.

80. JARA-BRAIN, Jülich-Aachen Research Alliance, Jülich, Germany.

81. Department of Neuroinformatics, Araya, Inc., Tokyo, Japan.

82. Sackler Centre for Consciousness Science, School of Psychology, University of Sussex, Falmer, UK.

83. Earth-Life Science Institute, Tokyo Institute of Technology, Tokyo, Japan.

84. Department of Psychiatry, McGill University, Montreal, QC, Canada.

85. McConnell Brain Imaging Center, Montreal Neurological Institute, Montreal, QC, Canada.

86. Department of Clinical and Experimental Epilepsy, UCL Queen Square Institute of Neurology, London, UK.

87. Genomics England, Queen Mary University of London, London, UK.

88. Public Psychiatry Division, Massachusetts Mental Health Center, Beth Israel Deaconess Medical Center, Harvard Medical School, Boston, MA, USA.

89. Psychiatry Neuroimaging Laboratory, Department of Psychiatry, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

90. Department of Genome Informatics, Graduate School of Medicine, Osaka University, Suita, Japan.

91. Department of Medical Biometry, Informatics and Epidemiology, University Hospital Bonn, Bonn, Germany.

92. Department of Psychology and Neuroscience, Duke University, Durham, NC, USA.

93. Section for Experimental Psychopathology and Neuroimaging, Department of General Psychiatry, Heidelberg University Hospital, Heidelberg, Germany.

94. Centre for Translational Research in Systems Neuroscience and Psychiatry, Department of Psychiatry and Psychotherapy, University Medical Center Göttingen, Göttingen, Germany.

95. Human Genetics, Genome Institute of Singapore, Singapore, Singapore.

96. MRC Centre for Neuropsychiatric Genetics and Genomics, Cardiff University, Cardiff, UK.

97. Department of Psychiatry, Harvard Medical School, Boston, MA, USA.

98. McGill Centre for Integrative Neuroscience, McGill University, Montreal, QC, Canada.

99. Department of Psychology, University of Edinburgh, Edinburgh, UK.

100. Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.

101. Department of Psychiatry and Human Behavior, School of Medicine University of California, Irvine, Irvine, CA, USA.

102. Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen, Netherlands.

103. Department of Psychiatry, Yale University School of Medicine, New Haven, CT, USA.

104. Olin Neuropsychiatric Research Center, Institute of Living, Hartford Hospital, Hartford, CT, USA.

105. Department of Medicine, University of Otago, Christchurch, Christchurch, New Zealand.

106. New Zealand Brain Research Institute, Christchurch, New Zealand.

107. Brain Research New Zealand - Rangahau Roro Aotearoa, Christchurch, New Zealand.

108. Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, Netherlands.

109. Institute of Neurogenomics, Helmholtz Zentrum München, German Research Centre for Environmental Health, Neuherberg, Germany.

110. IC - Institute of Computing, Campinas, Brazil.

111. Cécile and Oskar Vogt Institute of Brain Research, Medical Faculty, Heinrich Heine University, Düsseldorf, Germany.

112. Munich Cluster for Systems Neurology (SyNergy), Munich, Germany.

113. Institute of Translational Medicine, Liverpool, UK.

114. Drug Discovery Research, Astellas Pharmaceuticals, 21 Miyukigaoka, Tsukuba, Ibaraki 305-8585, Japan.

115. Department of Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, IN, USA.

116. Center for Neurobehavioral Genetics, University of California Los Angeles, Los Angeles, CA, USA.

117. NeuroSpin, CEA, Université Paris-Saclay, Gif-sur-Yvette, France.

118. Biostatistics and Computational Biology Unit, University of Otago, Christchurch, Christchurch, New Zealand.

119. Department of Pathology and Biomedical Science, University of Otago, Christchurch, Christchurch, New Zealand.

120. Department of Psychology, University of Oslo, Oslo, Norway.

121. Sunnaas Rehabilitation Hospital HT, Nesodden, Norway.

122. Department of Psychiatry, Radboud university medical center, Nijmegen, Netherlands.

123. Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, IN, USA.

124. Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.

125. Orygen, The National Centre of Excellence for Youth Mental Health, Melbourne, VIC, Australia.

126. The Centre for Youth Mental Health, University of Melbourne, Melbourne, VIC, Australia.

127. Department of Psychiatry, Vrije Universiteit University Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

128. School of Medical Sciences, University of New South Wales, Sydney, NSW, Australia.

129. Department of Biostatistics, Epidemiology and Informatics, University of Pennsylvania, Philadelphia, PA, USA.

130. Population Neuroscience & Developmental Neuroimaging, Bloorview Research Institute, University of Toronto, East York, ON, Canada.

131. Donders Centre for Cognitive Neuroimaging, Radboud University, Nijmegen, Netherlands.

132. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.

133. Centre for Healthy Brain Ageing, University of New South Wales, Sydney, NSW, Australia.

134. Neuroimaging Unit, Technological Facilities, Valdecilla Biomedical Research Institute IDIVAL, Santander, Spain.

135. Centro Investigacion Biomedica en Red Salud Mental, Santander, Spain.

136. Department of Psychology, Georgia State University, Atlanta, GA, USA.

137. Mind Research Network, Albuquerque, NM, USA.

138. Department of Psychiatry, University of Vermont, Burlington, VT, USA.

139. NORMENT, Division of Mental Health and Addiction, Oslo University Hospital and Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

140. School of Mental Health and Neuroscience, Faculty of Health, Medicine and Life Sciences, Maastricht University, Maastricht, Netherlands.

141. Max Planck Institute for Psycholinguistics, Nijmegen, Netherlands.

142. Department of Child and Adolescent Psychiatry/Psychology, Erasmus Medical Center-Sophia Children’s Hospital, Rotterdam, Netherlands.

143. Cognitive Neuroscience Center, Department of Biomedical Sciences of Cells and Systems, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.

144. Department of Neurology, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.

145. MRC Integrative Epidemiology Unit, Department of Population Health Sciences, Bristol Medical School, Bristol, UK.

146. Department of Psychology, University of Bath, Bath, UK.

147. UK Dementia Research Institute, The University of Edinburgh, Edinburgh, UK.

148. Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.

149. German Center for Neurodegenerative Diseases Rostock/Greifswald, Greifswald, Germany.

150. Department of Psychiatry, Psychosomatics and Psychotherapy, University of Würzburg, Würzburg, Germany.

151. Department of Neurology, FCM, UNICAMP, Campinas, Brazil.

152. Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK.

153. INSERM Unit 1000 - Neuroimaging & Psychiatry, Paris Saclay University, Gif sur Yvette, France.

154. Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria–IDIVAL, Santander, Spain.

155. Department of Psychiatry and Frank Porter Graham Child Development Institute, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.

156. Department of Psychiatry, University of California San Diego, San Diego, CA, USA.

157. VA San Diego Healthcare System, San Diego, CA, USA.

158. Avera Institute for Human Genetics, Sioux Falls, SD, USA.

159. Department of Psychiatry and Psychotherapy, Philipps-University Marburg, Marburg, Germany.

160. Department of Medical Genetics, Oslo University Hospital, Oslo, Norway.

161. NORMENT, Department of Clinical Science, University of Bergen, Bergen, Norway.

162. Department of Neurology, St James’s Hospital, Dublin, Ireland.

163. Academic Unit of Neurology, TBSI, Dublin, Ireland.

164. Future Neuro, Royal College of Surgeons in Ireland, Dublin, Ireland.

165. Information Sciences Institute, University of Southern California, Los Angeles, CA, USA.

166. Sir Peter Mansfield Imaging Centre, University of Nottingham, Nottingham, UK.

167. Brigham and Women’s Hospital, Boston, MA, USA.

168. The Broad Institute, Boston, MA, USA.

169. Harvard Medical School, Boston, MA, USA.

170. Center for Economics and Neuroscience, University of Bonn, Bonn, Germany.

171. Institute of Experimental Epileptology and Cognition Research, University Hospital Bonn, Germany.

172. Department of Clinical Radiology, University of Münster, Münster, Germany.

173. Department of Psychiatry, University of Iowa College of Medicine, Iowa City, IA, USA.

174. HMNC Holding GmbH, Munich, Germany.

175. University Medicine Greifswald, Interfaculty Institute for Genetics and Functional Genomics, Department of Functional Genomics, Greifswald, Germany.

176. Institute of Diagnostic Radiology and Neuroradiology, Greifswald, Germany.

177. Department of Radiology, Mayo Clinic, Rochester, MN, USA.

178. Core-Unit Brainimaging, Faculty of Medicine, University of Marburg, Marburg, Germany.

179. Department of Psychiatry and Behavioral Sciences, Duke University School of Medicine, Durham, NC, USA.

180. Department of Psychiatry, University of Minnesota, Minneapolis, MN, USA.

181. Department of Translational Neuroscience, UMC Utrecht Brain Center, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.

182. GGNet Mental Health, Apeldoorn, Netherlands.

183. Department of Psychiatry and Weill Institute for Neurosciences, University of California San Francisco, San Francisco, CA, USA.

184. Mental Health Service 116d, Veterans Affairs San Francisco Healthcare System, San Francisco, CA, USA.

185. Department of Neurology, Johns Hopkins University, Baltimore, MD, USA.

186. Department of Radiology, Johns Hopkins University, Baltimore, MD, USA.

187. Pacific Brain Health Center, Santa Monica, CA, USA.

188. John Wayne Cancer Institute, Santa Monica, CA, USA.

189. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.

190. German Centre for Cardiovascular Research, Greifswald, Germany.

191. Child and Adolescent Psychiatry Department, APHP Pitié Salpêtrière Hospital, Paris, France.

192. Radiology and Clinical Neurosciences, Hotchkiss Brain Institute, University of Calgary, Calgary, AB, Canada.

193. School of Medicine, University of California Irvine, Irvine, CA, USA.

194. School of Medicine and Public Health, University of Newcastle, Callaghan, NSW, Australia.

195. Priority Centre for Stroke and Brain Injury, University of Newcastle, Callaghan, NSW, Australia.

196. Hunter Medical Research Institute, Newcastle, NSW, Australia.

197. Department of Developmental Disability Neuropsychiatry, University of New South Wales, Sydney, NSW, Australia.

198. Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway.

199. School of Psychology, Trinity College Dublin, Dublin, Ireland.

200. Department of Radiology, Erasmus University Medical Centre, Rotterdam, Netherlands.

201. Department of Psychiatry, Osaka University Graduate School of Medicine, Suita, Japan.

202. Department of Epidemiology, Erasmus MC Medical Center, Rotterdam, Netherlands.

203. Department of Radiology and Nuclear Medicine, Erasmus MC Medical Center, Rotterdam, Netherlands.

204. Department of Clinical Genetics, Erasmus MC Medical Center, Rotterdam, Netherlands.

205. Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA.

206. INSERM, Bordeaux Population Health Research Center, team VINTAGE, UMR 1219, University of Bordeaux, Bordeaux, France.

207. Department of Neurology, CHU de Bordeaux, Bordeaux, France.

208. Department of Neurology, University of California, Davis, Sacramento, CA, USA.

209. Institute of Molecular Medicine, University of Texas Health Science Center at Houston, Houston, TX, USA.

210. Icelandic Heart Association, Kopavogur, Iceland.

211. Faculty of Medicine, University of Iceland, Reykjavik, Iceland.

212. Clinical Division of Neurogeriatrics, Department of Neurology, Medical University of Graz, Graz, Austria.

213. Institute for Medical Informatics, Statistics and Documentation, Medical University of Graz, Graz, Austria.

214. Laboratory of Epidemiology and Population Sciences, Intramural Research Program, National Institute on Aging, Bethesda, MD, USA.

215. Departments of Neurology and Epidemiology, University of Washington, Seattle, WA, USA.

216. Medical Informatics, Erasmus MC Medical Center, Rotterdam, Netherlands.

217. Neurodegeneratives Diseases Institute UMR 5293, CNRS, CEA, University of Bordeaux, Bordeaux, France.

218. MIND Center, University of Mississippi Medical Center, Jackson, MS, USA.

219. Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, University of Texas Health Sciences Center, San Antonio, TX, USA.

220. Department of Epidemiology & Biostatistics, University of Texas Health Sciences Center, San Antonio, TX, USA.

221. Department of Neurology, Boston University School of Medicine, Boston, MA, USA.

222. Framingham Heart Study and Department of Neurology, Boston University School of Medicine, Boston, MA, USA.

223. Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA.

224. Academic Unit for Psychiatry of Old Age, University of Melbourne, Melbourne, VIC, Australia.

225. National Ageing Research Institute, Melbourne, VIC, Australia.

226. Department of Neurology, Canterbury District Health Board, Christchurch, New Zealand.

227. Department of Psychiatry, Melbourne Medical School, University of Melbourne, Melbourne, VIC, Australia.

228. Florey Institute of Neuroscience and Mental Health, University of Melbourne, Parkville, VIC, Australia.

229. VA Mid-Atlantic Mental Illness Research Education and Clinical Center for Post Deployment Mental Health, Durham, VA Healthcare System, Durham, NC, USA.

230. Dementia Centre for Research Collaboration, University of New South Wales, Sydney, NSW, Australia.

231. Department of Clinical Genetics and School for Oncology and Developmental Biology (GROW), Maastricht University Medical Center, Maastricht, Netherlands.

232. Department of Psychology and Center for Brain Science, Harvard University, Boston, MA, USA.

233. Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.

234. Department of Radiology, Massachusetts General Hospital, Boston, MA, USA.

235. Karakter Child and Adolescent Psychiatry University Center, Nijmegen, Netherlands.

236. Department of Psychiatry, University of New Mexico, Albuquerque, NM, USA.

237. Department of Psychiatry, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.

238. Schizophrenia Research Institute, Randwick, NSW, Australia.

239. Tri-institutional Center for Translational Research in Neuroimaging and Data Science (TReNDS), Georgia State University, Georgia Institute of Technology, Emory University, Atlanta, GA, USA.

240. Department of Psychiatry, Monash University, Clayton, VIC, Australia.

241. Institute for Anatomy I, Medical Faculty, Heinrich-Heine University, Düsseldorf, Germany.

242. Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.

243. The SFI FutureNeuro Research Centre, Dublin, Ireland.

244. Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.

245. Hospital Universitario Virgen Del Rocio, IBiS, Universidad De Sevilla, Sevilla, Spain.

246. School of Psychology, Speech and Hearing, University of Canterbury, Christchurch, New Zealand.

247. Department of Neurology, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium.

248. Department of Cognitive and Clinical Neuroscience, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.

249. Department of Psychiatry (UPK), University of Basel, Basel, Switzerland.

250. Centre for Human Genetics, University of Marburg, Marburg, Germany.

251. Tommy Fuss Center for Neuropsychiatric Disease Research, Boston Children’s Hospital and Department of Psychiatry, Harvard Medical School, Boston, MA, USA.

252. Department of Neuroscience, Norwegian University of Science and Technology, Trondheim, Norway.

253. Department of Radiology and Nuclear Medicine, St. Olavs University Hospital, Trondheim, Norway.

254. University of Groningen, University Medical Center Groningen, Department of Psychiatry, Groningen, Netherlands.

255. Molecular Research Center for Children’s Mental Development, United Graduate School of Child Development, Osaka University, Suita, Japan.

256. Department of Pathology of Mental Diseases, National Institute of Mental Health, National Center of Neurology and Psychiatry, Tokyo, Japan.

257. Department of Psychiatry and Psychotherapy, Charité Campus Mitte, Charité - Universitätsmedizin Berlin, Berlin, Germany.

258. Health Behaviour Research Group, University of Newcastle, Callaghan, NSW, Australia.

259. Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands.

260. Department of Psychiatry, University Medical Center Groningen, University of Groningen, Groningen, Netherlands.

261. Department of Psychology, Yale University, New Haven, CT, USA.

262. Maryland Psychiatry Research Center, Department of Psychiatry, University of Maryland School of Medicine, Baltimore, MD, USA.

263. Department of Comparative Medicine, The University of Texas MD Anderson Cancer Center, Bastrop, TX, USA.

264. Center for Human Development, University of California San Diego, La Jolla, CA, USA.

265. Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, NY, USA.

266. Neurogenetics and Epigenetics, Brain and Mind Centre, The University of Sydney, Sydney, NSW, Australia.

267. Hunter New England Mental Health Service, Newcastle, NSW, Australia.

268. Herston Imaging Research Facility, School of Clinical Sciences, Queensland University of Technology, Brisbane, QLD, Australia.

269. Department of Psychiatry and Psychotherapy, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany.

270. School of Psychology, University of Newcastle, Callaghan, NSW, Australia.

271. Queensland Centre for Mental Health Research, University of Queensland, Brisbane, QLD, Australia.

272. Department of Radiation Sciences, Umeå University, Umeå, Sweden.

273. Emma Children's Hospital Academic Medical Center, Amsterdam, Netherlands.

274. Department of Pediatrics, Vrije Universiteit Medical Center, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

275. Clinical Neuropsychology section, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

276. NorthWestern Mental Health, Sunshine Hospital, St Albans, VIC, Australia.

277. Bloorview Research Institute, University of Toronto, Toronto, ON, Canada.

278. Departments of Psychology and Psychiatry, University of Toronto, Toronto, ON, Canada.

279. Centre for Developing Brain, Child Mind Institute, New York, NY, USA.

280. Department of Physiology, University of Toronto, Toronto, ON, Canada.

281. Department of Clinical Genetics, Vrije Universiteit Medical Centre, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.

282. Neuropsychiatric Institute, The Prince of Wales Hospital, Sydney, NSW, Australia.

283. PONS Research Group, Department of Psychiatry and Psychotherapie, Charité Campus Mitte, Humboldt University Berlin, Berlin, Germany.

284. Leibniz Institute for Neurobiology, Magdeburg, Germany.

285. Division of Molecular Medicine, John Hunter Hospital, New Lambton Heights, NSW, Australia.

286. General Psychiatry, Institute of Mental Health, Singapore, Singapore.

287. Chalfont Centre for Epilepsy, Chalfont-St-Peter, UK.

288. Stanley Center for Psychiatric Research, Broad Institute, Boston, MA, USA.

289. Department of Medical and Biological Psychology, University of Bergen, Bergen, Norway.

290. Department of Psychiatry and Neuroscience Institute, University of Cape Town, Cape Town, South Africa.

291. MRC Unit on Risk & Resilience in Mental Disorders, University of Cape Town, Cape Town, South Africa.

292. Department of Psychiatry, Leiden University Medical Center, Leiden, Netherlands.

293. Psychiatry, Neurology, Neuroscience, Genetics, Johns Hopkins University, Baltimore, MD, USA.

294. Centre for Advanced Imaging, University of Queensland, Brisbane, QLD, Australia.

295. Center for Cognitive Neuroscience, Neuroscience and Behavioral Disorders Program, Duke–National University of Singapore Medical School, Singapore, Singapore.

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 ( r G = −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

Funder

National Institutes of Health

National Institute of Mental Health

National Institute on Aging

U.S. National Library of Medicine

NIH Clinical Center

European Commission Directorate-General for Research and Innovation

National Health and Medical Research Council

NSW Health

UK Medical Research Council

Canadian Institutes of Health Research

National Natural Science Foundation of China