Article | Published:

Novel genetic loci underlying human intracranial volume identified through genome-wide association

Nature Neuroscience volume 19, pages 15691582 (2016) | Download Citation

Abstract

Intracranial volume reflects the maximally attained brain size during development, and remains stable with loss of tissue in late life. It is highly heritable, but the underlying genes remain largely undetermined. In a genome-wide association study of 32,438 adults, we discovered five previously unknown loci for intracranial volume and confirmed two known signals. Four of the loci were also associated with adult human stature, but these remained associated with intracranial volume after adjusting for height. We found a high genetic correlation with child head circumference (ρgenetic = 0.748), which indicates a similar genetic background and allowed us to identify four additional loci through meta-analysis (Ncombined = 37,345). Variants for intracranial volume were also related to childhood and adult cognitive function, and Parkinson's disease, and were enriched near genes involved in growth pathways, including PI3K-AKT signaling. These findings identify the biological underpinnings of intracranial volume and their link to physiological and pathological traits.

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References

  1. 1.

    , & Genetic variability of human brain size and cortical gyral patterns. Brain 120, 257–269 (1997).

  2. 2.

    & A new method for measuring cranial cavity volume and its application to the assessment of cerebral atrophy at autopsy. Neuropathol. Appl. Neurobiol. 3, 341–358 (1977).

  3. 3.

    , , , & Intracranial volume change in childhood. J. Neurosurg. 91, 610–616 (1999).

  4. 4.

    et al. A unified approach for morphometric and functional data analysis in young, old and demented adults using automated atlas-based head size normalization: reliability and validation against manual measurement of total intracranial volume. Neuroimage 23, 724–738 (2004).

  5. 5.

    et al. Maximal brain size remains an important predictor of cognition in old age, independent of current brain pathology. Neurobiol. Aging 33, 1758–1768 (2012).

  6. 6.

    et al. Common variants at 6q22 and 17q21 are associated with intracranial volume. Nat. Genet. 44, 539–544 (2012).

  7. 7.

    et al. Identification of common variants associated with human hippocampal and intracranial volumes. Nat. Genet. 44, 552–561 (2012).

  8. 8.

    et al. Common genetic variants influence human subcortical brain structures. Nature 520, 224–229 (2015).

  9. 9.

    et al. KCTD8 gene and brain growth in adverse intrauterine environment: a genome-wide association study. Cereb. Cortex 22, 2634–2642 (2012).

  10. 10.

    et al. Cohorts for Heart and Aging Research in Genomic Epidemiology (CHARGE) Consortium: design of prospective meta-analyses of genome-wide association studies from 5 cohorts. Circ Cardiovasc Genet 2, 73–80 (2009).

  11. 11.

    et al. The ENIGMA Consortium: large-scale collaborative analyses of neuroimaging and genetic data. Brain Imaging Behav. 8, 153–182 (2014).

  12. 12.

    et al. LD Score regression distinguishes confounding from polygenicity in genome-wide association studies. Nat. Genet. 47, 291–295 (2015).

  13. 13.

    et al. Defining the role of common variation in the genomic and biological architecture of adult human height. Nat. Genet. 46, 1173–1186 (2014).

  14. 14.

    et al. A common inversion under selection in Europeans. Nat. Genet. 37, 129–137 (2005).

  15. 15.

    & Tau pathology and neurodegeneration. Lancet Neurol. 12, 609–622 (2013).

  16. 16.

    et al. Genetic overlap between Alzheimer's disease and Parkinson's disease at the MAPT locus. Mol. Psychiatry 20, 1588–1595 (2015).

  17. 17.

    , , , & A 17q21.31 microduplication, reciprocal to the newly described 17q21.31 microdeletion, in a girl with severe psychomotor developmental delay and dysmorphic craniofacial features. Eur. J. Med. Genet. 50, 256–263 (2007).

  18. 18.

    et al. Mutations in the chromatin modifier gene KANSL1 cause the 17q21.31 microdeletion syndrome. Nat. Genet. 44, 639–641 (2012).

  19. 19.

    et al. Genome-wide meta-analysis identifies 56 bone mineral density loci and reveals 14 loci associated with risk of fracture. Nat. Genet. 44, 491–501 (2012).

  20. 20.

    et al. Phenotypic dissection of bone mineral density reveals skeletal site specificity and facilitates the identification of novel loci in the genetic regulation of bone mass attainment. PLoS Genet. 10, e1004423 (2014).

  21. 21.

    et al. GWAS of longevity in CHARGE consortium confirms APOE and FOXO3 candidacy. J. Gerontol. A Biol. Sci. Med. Sci. 70, 110–118 (2015).

  22. 22.

    et al. A genome-wide association study identifies novel loci associated with circulating IGF-I and IGFBP-3. Hum. Mol. Genet. 20, 1241–1251 (2011).

  23. 23.

    et al. FoxOs cooperatively regulate diverse pathways governing neural stem cell homeostasis. Cell Stem Cell 5, 540–553 (2009).

  24. 24.

    et al. FoxO3 regulates neural stem cell homeostasis. Cell Stem Cell 5, 527–539 (2009).

  25. 25.

    et al. Associations of IGF-1 gene variants and milk protein intake with IGF-I concentrations in infants at age 6 months: results from a randomized clinical trial. Growth Horm. IGF Res. 23, 149–158 (2013).

  26. 26.

    et al. Common variants at 12q15 and 12q24 are associated with infant head circumference. Nat. Genet. 44, 532–538 (2012).

  27. 27.

    et al. The 12q14 microdeletion syndrome: six new cases confirming the role of HMGA2 in growth. Eur. J. Hum. Genet. 19, 534–539 (2011).

  28. 28.

    , , & Evidence for involvement of NFBP in processing of ribosomal RNA. J. Cell. Physiol. 214, 381–388 (2008).

  29. 29.

    et al. Multi-ethnic genome-wide association study of cerebral white matter hyperintensities on MRI. Circgenetics. 114, 000858 (2015).

  30. 30.

    , , , & GEMC1 is a TopBP1-interacting protein required for chromosomal DNA replication. Nat. Cell Biol. 12, 484–491 (2010).

  31. 31.

    et al. Genome-wide analysis of body proportion classifies height-associated variants by mechanism of action and implicates genes important for skeletal development. Am. J. Hum. Genet. 96, 695–708 (2015).

  32. 32.

    et al. Head circumference and body growth in autism spectrum disorders. Brain Dev. 33, 569–575 (2011).

  33. 33.

    Cognitive reserve in ageing and Alzheimer's disease. Lancet Neurol. 11, 1006–1012 (2012).

  34. 34.

    , & PI3 kinase/Akt activation mediates estrogen and IGF-1 nigral DA neuronal neuroprotection against a unilateral rat model of Parkinson's disease. Dev. Neurobiol. 68, 632–644 (2008).

  35. 35.

    Brain overgrowth in disorders of RTK-PI3K-AKT signaling: a mosaic of malformations. Semin. Perinatol. 39, 36–43 (2015).

  36. 36.

    et al. De novo germline and postzygotic mutations in AKT3, PIK3R2 and PIK3CA cause a spectrum of related megalencephaly syndromes. Nat. Genet. 44, 934–940 (2012).

  37. 37.

    et al. Mapping of deletion and translocation breakpoints in 1q44 implicates the serine/threonine kinase AKT3 in postnatal microcephaly and agenesis of the corpus callosum. Am. J. Hum. Genet. 81, 292–303 (2007).

  38. 38.

    , , & Duplication of AKT3 as a cause of macrocephaly in duplication 1q43q44. Am. J. Med. Genet. A. 161A, 2016–2019 (2013).

  39. 39.

    et al. Association of common genetic variation in the insulin/IGF1 signaling pathway with human longevity. Aging Cell 8, 460–472 (2009).

  40. 40.

    et al. Genetic contributions to variation in general cognitive function: a meta-analysis of genome-wide association studies in the CHARGE consortium (N = 53,949). Mol. Psychiatry 20, 183–192 (2015).

  41. 41.

    et al. Meta-analysis of genome-wide association studies for neuroticism, and the polygenic association with major depressive disorder. JAMA Psychiatry 72, 642–650 (2015).

  42. 42.

    et al. Meta-analysis of genome-wide association studies for extraversion: findings from the Genetics of Personality Consortium. Behav. Genet. 46, 170–182 (2016).

  43. 43.

    et al. Large-scale meta-analysis of genome-wide association data identifies six new risk loci for Parkinson's disease. Nat. Genet. 46, 989–993 (2014).

  44. 44.

    Cross-Disorder Group of the Psychiatric Genomics Consortium. Identification of risk loci with shared effects on five major psychiatric disorders: a genome-wide analysis. Lancet 381, 1371–1379 (2013).

  45. 45.

    et al. Childhood intelligence is heritable, highly polygenic and associated with FNBP1L. Mol. Psychiatry 19, 253–258 (2014).

  46. 46.

    et al. Integrative analysis of 111 reference human epigenomes. Nature 518, 317–330 (2015).

  47. 47.

    , & METAL: fast and efficient meta-analysis of genomewide association scans. Bioinformatics 26, 2190–2191 (2010).

  48. 48.

    et al. Association of Alzheimer's disease GWAS loci with MRI markers of brain aging. Neurobiol. Aging 36, 1765.e7–1765.e16 (2015).

  49. 49.

    et al. Partitioning heritability by functional category using GWAS summary statistics. Nat. Genet. 47, 1228–1235 (2015).

  50. 50.

    et al. New loci associated with birth weight identify genetic links between intrauterine growth and adult height and metabolism. Nat. Genet. 45, 76–82 (2013).

  51. 51.

    et al. A novel common variant in DCST2 is associated with length in early life and height in adulthood. Hum. Mol. Genet. 24, 1155–1168 (2015).

  52. 52.

    et al. Meta-analysis of 74,046 individuals identifies 11 new susceptibility loci for Alzheimer's disease. Nat. Genet. 45, 1452–1458 (2013).

  53. 53.

    , , & GATES: a rapid and powerful gene-based association test using extended Simes procedure. Am. J. Hum. Genet. 88, 283–293 (2011).

  54. 54.

    , , , & Common inherited variation in mitochondrial genes is not enriched for associations with type 2 diabetes or related glycemic traits. PLoS Genet. 6, e1001058 (2010).

  55. 55.

    , & HYST: a hybrid set-based test for genome-wide association studies, with application to protein-protein interaction-based association analysis. Am. J. Hum. Genet. 91, 478–488 (2012).

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Acknowledgements

CHARGE: Infrastructure for the CHARGE Consortium is supported in part by the National Heart, Lung and Blood Institute grant HL105756 and for the neuroCHARGE phenotype working group through the National Institute on Aging grant AG033193.

ENIGMA: ENIGMA was supported in part by a Consortium grant (U54 EB020403 to PMT) from the NIH Institutes contributing to the Big Data to Knowledge (BD2K) Initiative, including the NIBIB and NCI.

Age, Gene/Environment Susceptibility-Reykjavik Study (AGES-Reykjavik): This study has been funded by NIH contracts N01-AG-1-2100 and 271201200022C, the NIA Intramural Research Program, Hjartavernd (the Icelandic Heart Association), and the Althingi (the Icelandic Parliament). The study was approved by the Icelandic National Bioethics Committee, VSN: 00-063. The researchers are indebted to the participants for their willingness to participate in the study.

Alzheimer's Disease Neuroimaging Initiative (ADNI): Data collection and sharing for this project was funded by the ADNI (National Institutes of Health Grant U01 AG024904) and DOD ADNI (Department of Defense award number W81XWH-12-2-0012). ADNI is funded by the National Institute on Aging, the National Institute of Biomedical Imaging and Bioengineering, and through generous contributions from the Alzheimer's Association; Alzheimer's Drug Discovery Foundation; BioClinica, Inc.; Biogen Idec Inc.; Bristol-Myers Squibb Company; Eisai Inc.; Elan Pharmaceuticals, Inc.; Eli Lilly and Company; F. Hoffmann-La Roche Ltd and its affiliated company Genentech, Inc.; GE Healthcare; Innogenetics, N.V.; IXICO Ltd.; Janssen Alzheimer Immunotherapy Research & Development, LLC.; Johnson & Johnson Pharmaceutical Research & Development LLC.; Medpace, Inc.; Merck & Co., Inc.; Meso Scale Diagnostics, LLC.; NeuroRx Research; Novartis Pharmaceuticals Corporation; Pfizer Inc.; Piramal Imaging; Servier; Synarc Inc.; and Takeda Pharmaceutical Company. The Canadian Institutes of Health Research provides funds to support ADNI clinical sites in Canada. Private sector contributions are facilitated by the Foundation for the National Institutes of Health (http://www.fnih.org). The grantee organization is the Northern California Institute for Research and Education, and the study is coordinated by the Alzheimer's Disease Cooperative Study at the University of California, San Diego. ADNI data are disseminated by the Laboratory of Neuro Imaging at the University of Southern California.

ANM: AddNeuroMed was funded through the EU FP6 programme. HS: Academy of Finland, Research Council for Health, 258081, UEFBrain, University of Eastern Finland, VTR funding Kuopio University Hospital.

Atherosclerosis Risk In Communities Study (ARIC): The Atherosclerosis Risk in Communities study was performed as a collaborative study supported by National Heart, Lung, and Blood Institute (NHLBI) contracts (HHSN268201100005C, HSN268201100006C, HSN268201100007C, HHSN268201100008C, HHSN268201100009C, HHSN268201100010C, HHSN268201100011C, and HHSN268201100012C), R01HL70825, R01HL087641, R01HL59367, and R01HL086694; National Human Genome Research Institute contract U01HG004402; and National Institutes of Health (NIH) contract HHSN268200625226C. Infrastructure was partly supported by grant no. UL1RR025005, a component of the NIH and NIH Roadmap for Medical Research. This project was also supported by NIH R01 grant NS087541 to M.F.

Austrian Stroke Prevention Study Family (ASPS-Fam): The ASPS-Fam is funded by the Austrian Science Fund (FWF) project I904, the Medical University of Graz and the Steiermärkische Krankenanstalten Gesellschaft.

BETULA: This sample collection was supported by a Wallenberg Scholar grant from the Knut and Alice Wallenberg (KAW) foundation and a grant from Torsten and Ragnar Söderbergs Foundation to Lars Nyberg. S.l.H. was supported by a grant from HelseVest RHF (grant 911554).

Bipolar Family Study (BFS): The Bipolar Family Study wishes to thank the Scottish Mental Health Research Network for research assistant support, the Brain Research Imaging Centre Edinburgh, a center in the Scottish Funding Council Scottish Imaging Network–A Platform for Scientific Excellence (SINAPSE) Collaboration, for image acquisition and the Wellcome Trust Clinical Research Facility for genotyping. Genotyping was supported by the National Alliance for Research on Schizophrenia and Depression (NARSAD) Independent Investigator Award (to A.M.M.), and data collection was supported by the Health Foundation Clinician Scientist Fellowship. The research leading to these results also receives funding from the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements #602450 (IMAGEMEND) and ongoing support from the Wellcome Trust (Ref 104036/Z/14/Z).

Brain Imaging Genetics (BIG): This work makes use of the BIG database, first established in Nijmegen, The Netherlands, in 2007. This resource is now part of Cognomics (http://www.cognomics.nl), 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 in Nijmegen. The Board of the Cognomics Initiative consists of B. Franke, S. Fisher, G. Fernandez, P. Hagoort, H. Brunner, J. Buitelaar, H. van Bokhoven and D. Norris. The Cognomics Initiative has received supported from the participating departments and centers and from external grants, that is, the Biobanking and Biomolecular Resources Research Infrastructure (Netherlands) (BBMRI-NL), the Hersenstichting Nederland, and the Netherlands Organization for Scientific Research (NWO). The research leading to these results also receives funding from the NWO Gravitation grant 'Language in Interaction', the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements no 602450 (IMAGEMEND), no 278948 (TACTICS), and no 602805 (Aggressotype), as well as from the European Community's Horizon 2020 programme under grant agreement no 643051 (MiND) and from ERC-2010-AdG 268800-NEUROSCHEMA. In addition, the work was supported by a grant for the ENIGMA Consortium (grant number U54 EB020403) from the BD2K Initiative of a cross-NIH partnership. We wish to thank all persons who kindly participated in the BIG research.

Brain Genomics Superstruct Project (GSP): Data were provided in part by the Brain Genomics Superstruct Project of Harvard University and the Massachusetts General Hospital, with support from the Center for Brain Science Neuroinformatics Research Group, the Athinoula A. Martinos Center for Biomedical Imaging, and the Center for Human Genetic Research. 20 individual investigators at Harvard and MGH generously contributed data to GSP. This work is supported by NIMH grants K99 MH101367 (P.H.L.), R01-MH079799 (J.W.S.), K24MH094614 (J.W.S.) and K01MH099232 (A.J.H.).

Brainscale and NTR-Adults: We would like to thank all twin participants from the Netherlands Twin Register. The NTR-adult and Brainscale studies were supported by the Netherlands Organization for Scientific Research NWO [MW904-61-193 (E.d.G. and D.B.), MaGW-nr: 400-07-080 (D. V't E.), MagW 480-04-004 (D.B.), (51.02.060 (H.H.), 668.772 (D.B. and H.H.); NWO/SPI 56-464-14192 (D.B.), the European Research Council (ERC-230374) (D.B.), High Potential Grant Utrecht University (H.H.), NWO Brain and Cognition 433-09-220 (H.H.) and the Neuroscience Campus Amsterdam.

Cardiovascular Health Study (CHS): This research was supported by NHLBI contracts HHSN268201200036C, HHSN268200800007C, N01HC55222, N01HC85079, N01HC85080, N01HC85081, N01HC85082, N01HC85083, N01HC85086; and NHLBI grants U01HL080295, R01HL087652, R01HL105756, R01HL103612, R01HL120393, and R01HL130114 with additional contribution from the National Institute of Neurological Disorders and Stroke (NINDS). Additional support was provided through R01AG023629, R01AG15928, R01AG20098, R01AG027002, R01AG05133, and R01AG027058 from the National Institute on Aging (NIA). A full list of principal CHS investigators and institutions can be found at http://CHS-NHLBI.org. 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 (DRC) grant DK063491 to the Southern California Diabetes Endocrinology Research Center. The content is solely the responsibility of the authors and does not necessarily represent the official views of the National Institutes of Health.

CHAP: This research was funded by grants from the National Institute of Health (AG011101 and AG030146) and the International Alzheimer's Association (NIRP-14-302587). DNA samples were collected during clinical evaluations and population interviews, and analyzed at the Broad Institute.

Epidemiology of Dementia in Singapore (EDIS): The Singapore Malay Eye Study (SiMES) and the Singapore Chinese Eye. Study (SCES) are funded by National Medical Research Council (grants 0796/2003, IRG07nov013, IRG09nov014, STaR/0003/2008 and CG/SERI/2010) and Biomedical Research Council (grants 09/1/35/19/616), Singapore. The Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore provided services for genotyping. The Epidemiology of Dementia in Singapore study is supported by the National Medical Research Council, Singapore (NMRC/CG/NUHS/2010, grant no. R-184-006-184-511). M.K.I. received additional funding from the Singapore Ministry of Health's National Medical Research Council (NMRC/CSA/038/2013).

EPIGEN: Work from the London Cohort was supported by research grants from the Wellcome Trust (grant 084730 to S.M.S.), University College London (UCL)/University College London Hospitals (UCLH) NIHR Biomedical Research Centre/Specialist Biomedical Research Centres (CBRC/SBRC) (grant 114 to S.M.S.), the European Union Marie Curie Reintegration (to M. Matarin and S.M.S.), the UK NIHR (08-08-SCC), the Comprehensive Local Research Network (CLRN) Flexibility and Sustainability Funding (FSF) (grant CEL1300 to S.M.S.), The Big Lottery Fund, the Wolfson Trust and the Epilepsy Society. This work was undertaken at UCLH/UCL, which received a proportion of funding from the UK Department of Health's NIHR Biomedical Research Centres funding scheme. Work from the Royal College of Surgeons in Ireland was supported by research grants from the Science Foundation Ireland (Research Frontiers Programme award 08/RFP/GEN1538) and Brainwave–the Irish Epilepsy Association. M. Matarin is funded by Epilepsy Research UK (grant F1206).

Erasmus Rucphen Family study (ERF) The ERF study as a part of EUROSPAN (European Special Populations Research Network) was supported by European Commission FP6 STRP grant number 018947 (LSHG-CT-2006-01947) and also received funding from the European Community's Seventh Framework Programme (FP7/2007-2013)/grant agreement HEALTH-F4-2007-201413 by the European Commission under the programme “Quality of Life and Management of the Living Resources” of 5th Framework Programme (no. QLG2-CT-2002-01254). High-throughput analysis of the ERF data was supported by joint grant from Netherlands Organization for Scientific Research and the Russian Foundation for Basic Research (NWO-RFBR 047.017.043). We are grateful to all study participants and their relatives, general practitioners and neurologists for their contributions and to P. Veraart for her help in genealogy, J. Vergeer for the supervision of the laboratory work and P. Snijders for his help in data collection. N. Amin is supported by the Netherlands Brain Foundation (project number F2013(1)-28). The ERF study genome-wide array data and phenotype data (age and gender) is archived in European Genome-Phenome Database (EGA). The study is archived in the DAC named Erasmus Rucphen Family Study with the accession code: EGAS00001001134. Researchers who wish to use other phenotypic data of the Erasmus Rucphen Family Study must seek approval from the management team of the Erasmus Rucphen Family study. They are advised to contact Cornelia van Duijn (c.vanduijn@erasmusmc.nl).

Framingham Heart Study (FHS): This work was supported by the dedication of the Framingham Study participants, the National Heart, Lung and Blood Institute's Framingham Heart Study (Contract No. HHSN268201500001I), and by grants from the National Institute of Health (AG008122, AG054076, AG049607, AG033193, AG010129, NS017950, and U01AG49505).

Generation R: The Generation R Study is conducted by the Erasmus Medical Centre in close collaboration with the Municipal Health Service Rotterdam area, and the Stichting Trombosedienst & Artsenlaboratorium Rijnmond (STAR), Rotterdam. We gratefully acknowledge the contribution of general practitioners, hospitals, midwives and pharmacies in Rotterdam. Additional support for neuroimaging came from ZonMw TOP 40-00812-98-11021.

GeneSTAR: GeneSTAR was supported by grants from the National Institute of Neurological Disorders and Stroke (R01NS062059), the National Heart, Lung, and Blood Institute (U01 HL72518, HL097698) and the National Institutes of Health/National Center for Research Resources (M01-RR000052) to the Johns Hopkins General Clinical Research Center. We would like to thank the participants and families of GeneSTAR and our dedicated staff for all their sacrifices.

GIG: The GIG (Genomic Imaging Göttingen) sample was established at the Center for Translational Research in Systems Neuroscience and Psychiatry at Göttingen University. We thank M. Keil, E. Diekhof, T. Melcher and I. Henseler for assistance in MRI data acquisition, and E. Binder and H. Mohr for their valuable help with genotyping. We are grateful to all persons who kindly participated in the GIG study.

GOBS: We acknowledge the ultimate source of our data, the Mexican American community of San Antonio and surrounding areas. Financial support for this study was provided by grants from the National Institute of Mental Health MH0708143 (D.C. Glahn), MH078111 (J. Blangero) and MH083824 (D.C. Glahn and J. Blangero). Theoretical development of SOLAR is supported by MH59490 (J. Blangero). This investigation was conducted, in part, in facilities constructed with support from Research Facilities Improvement Program grant numbers C06 RR13556 and C06 RR017515 from the National Center for Research Resources, NIH. Some of this work was performed at Texas Biomedical Research Institute, where J. Blangero began this investigator-initiated competitively publicly funded work.

HUBIN: This study was financed by the Swedish Research Council (K2007-62X-15077-04-1, K2008-62P-20597-01-3. K2010-62X-15078-07-2, K2012-61X-15078-09-3), 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. Genotyping was performed by the SNP&SEQ Technology Platform in Uppsala. The platform is part of Science for Life Laboratory at Uppsala University and supported as a national infrastructure by the Swedish Research Council.

IMAGEN: IMAGEN was supported by the European Union-funded FP6 Integrated Project IMAGEN (Reinforcement-related behavior in normal brain function and psychopathology) (LSHM-CT- 2007-037286), the FP7 projects IMAGEMEND (602450) and MATRICS (603016), and the Innovative Medicine Initiative Project EU-AIMS (115300-2), the Medical Research Council Programme Grant “Developmental pathways into adolescent substance abuse” (93558), as well as the NIHR-biomedical Research Center “Mental Health”. Further support was provided by the Swedish Research Council FORMAS, and the German Federal Ministry for Education and Research BMBF (eMED SysAlc 01ZX1311A; Forschungsnetz AERIAL; 1EV0711) and the US National Institutes of Health (Axon, Testosterone and Mental Health during Adolescence; MH085772-01A1), and grants from the French MILDECA and from the Fondation pour la Recherche Médicale.

IMpACT: This study was funded by a grant from the Brain & Cognition Excellence Program of the Netherlands Organization for Scientific Research (NWO, grant 433-09-229) and in part by the Netherlands Brain Foundation (grant number, 15F07[2]27). B. Franke is supported by a Vici grant from the Netherlands Organization for Scientific Research (NWO; grant no 016.130.669). The research leading to these results also receives funding from the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements no 602450 (IMAGEMEND), no 278948 (TACTICS), and no 602805 (Aggressotype) as well as from the European Community's Horizon 2020 programme under grant agreement no 643051 (MiND). In addition, the work was supported by a grant for the ENIGMA Consortium (grant number U54 EB020403) from the BD2K Initiative of a cross-NIH partnership.

LBC1936: We thank the LBC1936 participants and the members of the LBC1936 research team who collected and collated the phenotypic and genotypic data. This work was undertaken as part of the Cross Council and University of Edinburgh Centre for Cognitive Ageing and Cognitive Epidemiology (CCACE; http://www.ccace.ed.ac.uk). This work was supported by a Research into Ageing programme grant (to I.J.D.) and the Age UK-funded Disconnected Mind project (http://www.disconnectedmind.ed.ac.uk; to I.J.D. and J.M.W.), with additional funding from the UK Medical Research Council (MRC; to I.J.D., J.M.W. and M.E.B.). J.M.W. is supported by the Scottish Funding Council through the SINAPSE Collaboration (http://www.sinapse.ac.uk). M.V.M. is supported by the Row Fogo Charitable Trust. CCACE (MRC MR/K026992/1) is funded by the UK Biotechnology and Biological Sciences Research Council (BBSRC) and the UK MRC. Genotyping was supported by a grant from the BBSRC (BB/F019394/1).The image acquisition and analysis was performed at the Brain Research Imaging Centre, University of Edinburgh (http://www.bric.ed.ac.uk).

Leiden Longevity Study (LLS): The Leiden Longevity Study was supported by a grant from the Innovation-Oriented Research Program on Genomics (SenterNovem IGE05007) and the Netherlands Consortium for Healthy Ageing (grant number 050-060-810).

Mind Clinical Imaging Consortium (MCIC): Data used in the preparation of this work were obtained from the Mind Clinical Imaging Consortium database through the Mind Research Network (http://www.mrn.org). The MCIC project was supported by the Department of Energy under Award Number DE-FG02-08ER64581. MCIC is the result of efforts of co-investigators from University of Iowa, University of Minnesota, University of New Mexico, and Massachusetts General Hospital.

MooDS: The establishment of the MooDS sample was funded by the German Federal Ministry of Education and Research (BMBF) through the Integrated Genome Research Network (IG) MooDS (Systematic Investigation of the Molecular Causes of Major Mood Disorders and Schizophrenia; grant 01GS08144 to M.M. Nöthen and S. Cichon, grant 01GS08147 to J. Rietschel and A. Meyer-Lindenberg and grant 01GS08148 to A. Heinz), under the auspices of the National Genome Research Network plus (NGFNplus), and through the Integrated Network IntegraMent (Integrated Understanding of Causes and Mechanisms in Mental Disorders), under the auspices of the e:Med Programme (grant 01ZX1314A to M.M. Nöthen, grant 01ZX1314C to H. Walter, grant 01ZX1314G to M. Rietschel).

MPIP: The MPIP Munich Morphometry Sample comprises images acquired as part of the Munich Antidepressant Response Signature Study and the Recurrent Unipolar Depression (RUD) Case-Control study performed at the MPIP, and control subjects acquired at the Ludwig-Maximilians-University, Munich, Department of Psychiatry. We thank E. Meisenzahl and D. Rujescu for providing MRI and genetic data for inclusion into the MPIP Munich Morphometry sample. We wish to acknowledge A. Olynyik and radiographers R. Schirmer, E. Schreiter, R. Borschke and I. Eidner for image acquisition and data preparation. We thank D.P. Auer for local study management in the initial phase of the RUD study. We are grateful to GlaxoSmithKline for providing the genotypes of the Recurrent Unipolar Depression Case-Control Sample. We thank the staff of the Center of Applied Genotyping (CAGT) for generating the genotypes of the MARS cohort. The study is supported by a grant of the Exzellenz-Stiftung of the Max Planck Society. This work has also been funded by the Federal Ministry of Education and Research (BMBF) in the framework of the National Genome Research Network (NGFN), FKZ 01GS0481.

NCNG: this sample collection was supported by grants from the Bergen Research Foundation and the University of Bergen, the Dr. Einar Martens Fund, the K.G. Jebsen Foundation, the Research Council of Norway, to S.L.H., V.M.S. and T.E.

NESDA: Funding was obtained from the Netherlands Organization for Scientific Research (Geestkracht program grant 10-000-1002); the Center for Medical Systems Biology (CSMB, NWO Genomics), Biobanking and Biomolecular Resources Research Infrastructure (BBMRI-NL), VU University's Institutes for Health and Care Research (EMGO+) and Neuroscience Campus Amsterdam, University Medical Center Groningen, Leiden University Medical Center, National Institutes of Health (NIH, R01D0042157-01A, MH081802, Grand Opportunity grants 1RC2 MH089951 and 1RC2 MH089995). Part of the genotyping and analyses were funded by the Genetic Association Information Network (GAIN) of the Foundation for the National Institutes of Health. Computing was supported by BiG Grid, the Dutch e-Science Grid, which is financially supported by NWO.

NeuroIMAGE: The NeuroIMAGE project was supported by NIH Grant R01MH62873 (to S.V. Faraone), NWO Large Investment Grant 1750102007010 (to J. Buitelaar), and by grants from Radboud University Nijmegen Medical Center, University Medical Center Groningen and Accare, and VU University Amsterdam. The work contributing to this result also receives support from the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements no 602450 (IMAGEMEND), no 278948 (TACTICS) and no 602805 (Aggressotype) as well as from the European Community's Horizon 2020 programme under grant agreement n0 643051 (MiND). In addition, the work was supported by a grant for the ENIGMA Consortium (grant number U54 EB020403) from the BD2K Initiative of a cross-NIH partnership.

NIMH-IRP: Supported in part by the NIMH Intramural Research Program (ZIAMH002810; Z01MH002792; Z01MH002790).

North American Brain Expression Consortium (NABEC): This research was supported by the Intramural Research Program of the NIH, National Institute on Aging.

Older Australian Twins Study (OATS): We would like to acknowledge and thank the OATS participants, their supporters and respective Research Teams. This work was supported by a number of sources. OATS is supported by the NHMRC/Australian Research Council Strategic Award 401162 and NHMRC Project Grant 1045325 to P. Sachdev and colleagues. OATS was facilitated through access to the Australian Twin Registry, a national research resource supported by the NHMRC Enabling Grant 310667, administered by the University of Melbourne. DNA was extracted by Genetic Repositories Australia, an Enabling Facility supported by the NHMRC Grant 401184. OATS genotyping was partly funded by a Commonwealth Scientific and Industrial Research Organisation Flagship Collaboration Fund Grant. Henry Brodaty is supported by the Australian Government funded Dementia Collaborative Research Centre (DCRC), UNSW. Nicola Armstrong was supported by the NHMRC Project Grant 525453 and Karen Mather is supported by an Alzheimer's Australia Dementia Research Foundation Postdoctoral Fellowship and the NHMRC Capacity Building Grant 568940.

Osaka: This study was supported, in part, by research grants from the Japanese Ministry of Health, Labor and Welfare; the Japanese Ministry of Education, Culture, Sports, Science and Technology (MEXT) KAKENHI and Scientific Research on Innovative Areas (Comprehensive Brain Science Network); and the Brain Sciences Project of the Center for Novel Science Initiatives (CNSI), the National Institutes of Natural Sciences (NINS), and the Brain Mapping by Integrated Neurotechnologies for Disease Studies (Brain/MINDS) from Japan Agency for Medical Research and development, AMED.

PAFIP: The PAFIP study was supported by Instituto de Salud Carlos III, FIS00/3095, 01/3129, PI020499, PI060507, PI10/00183, PI14/00639, the SENY Fundació Research Grant CI 2005-0308007, and the Fundación Marqués de Valdecilla API07/011. PAFIP wish to acknowledge WTCCC2 (Wellcome Trust Case Control Consortium 2) for DNA Genotyping, Valdecilla Biobank for providing the biological samples and associated data included in this study and Idival Neuroimaging Unit for its help in the technical execution of this work. D. Tordesillas-Gutiérrez is funded by a contract from the Carlos III Health Institute (CA12/00312).

PROSPER: The PROSPER study was supported by an investigator-initiated grant obtained from Bristol-Myers Squibb. J.W. Jukema is an Established Clinical Investigator of the Netherlands Heart Foundation (grant 2001 D 032). Support for genotyping was provided by the seventh framework program of the European commission (grant 223004) and by the Netherlands Genomics Initiative (Netherlands Consortium for Healthy Aging grant 050-060-810).

QTIM: D.P.H., N.J., C.R.K.C. and P.M.T. are supported, in part, by NIH grants R01 NS080655, R01AG040060, R01 EB008432, R01 MH097268, U01 AG024904, R01 MH085667, R01 MH089722, P41 EB015922, and R01 MH094343. R.K.W. is supported by National Science Foundation (BCS-1229450). J.L.S. was supported by the NIMH (K99MH102357) and Autism Speaks. G.Z. is supported by a Future Fellowship (FT0991634) from the Australian Research Council. S.E.M. and G.W.M. are supported by a National Health and Medical Research Council (NHMRC), Australia, Fellowships (1103623, 619667). The QTIM study is supported by grants from NIH (R01 HD050735) and the NHMRC (389875, 486682, 1009064). We thank the twins and siblings for their participation, M. Grace and A. Eldridge for twin recruitment, A. Al Najjar and other radiographers for scanning, K. McAloney and D. Park for research support, and A. Henders and staff for DNA sample processing and preparation.

ROS and MAP: The clinical, genomic, and neuroimaging data for the Religious Orders Study and the Rush Memory and Aging Project was funded by NIH grants P30AG10161, RF1AG15819, R01AG17917, R01AG30146, R01AG40039, and the Translational Genomics Research Institute.

Rotterdam Study: The generation and management of GWAS genotype data for the Rotterdam Study are supported by the Netherlands Organisation of Scientific Research NWO Investments (nr. 175.010.2005.011, 911-03-012). This study is funded by the Research Institute for Diseases in the Elderly (014-93-015; RIDE2), the Netherlands Genomics Initiative (NGI)/Netherlands Organisation for Scientific Research (NWO) project nr. 050-060-810. The Rotterdam Study is funded by Erasmus Medical Center and Erasmus University, Rotterdam, Netherlands Organization for the Health Research and Development (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. This research is supported by the Dutch Technology Foundation STW, which is part of the NWO, and which is partly funded by the Ministry of Economic Affairs. This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement No 678543). Further support was obtained through the Joint Programme - Neurodegenerative Disease Research working group on High-Dimensional Research in Alzheimer's Disease (ZonMW grant number 733051031). MAI is supported by ZonMW grant number 916.13.054. H.H.H.A. is supported by the Van Leersum Grant of the Royal Netherlands Academy of Arts and Sciences.

Saguenay Youth Study (SYS): The Saguenay Youth Study project is funded by the Canadian Institutes of Health Research (T.P., Z.P.), Heart and Stroke Foundation of Quebec (Z.P.), and the Canadian Foundation for Innovation (Z.P.). T.P. is supported by the Tanenbaum Chair in Population Neuroscience at the Rotman Research Institute, University of Toronto.

SHIP and TREND: The SHIP data sets are part of the Community Medicine Research net (CMR) of the University of Greifswald, which is funded by the German Federal Ministry of Education and Research and the German Ministry of Cultural Affairs, as well as by the Social Ministry of the Federal State of Mecklenburg–West Pomerania (grants no. 01ZZ9603, 01ZZ0103, and 01ZZ0403), and the network 'Greifswald Approach to Individualized Medicine (GANI_MED)' funded by the Federal Ministry of Education and Research (grant 03IS2061A). Genome-wide data and MRI scans were supported by the Federal Ministry of Education and Research (grant no. 03ZIK012) and a joint grant from Siemens Healthcare, Erlangen, Germany, and the Federal State of Mecklenburg–West Pomerania. The University of Greifswald is a member of t the Caché Campus Program of the InterSystems GmbH.

Sydney Memory and Ageing Study (Sydney MAS): We would like to thank the Sydney MAS participants, their supporters and respective research teams. Sydney MAS was supported by the Australian National Health and Medical Research Council (NHMRC) Program Grants 350833 and 568969 to P. Sachdev, H. Brodaty and G. Andrews. DNA was extracted by Genetic Repositories Australia, an Enabling Facility supported by the NHMRC Grant 401184. H. Brodaty is supported by the Australian Government funded Dementia Collaborative Research Centre (DCRC), UNSW. N. Armstrong was supported by the NHMRC Project Grant 525453 and K. Mather is supported by an Alzheimer's Australia Dementia Research Foundation Postdoctoral Fellowship. Both S. Reppermund and K. Mather are supported by the NHMRC Capacity Building Grant 568940.

Tasmanian Study of Gait and Cognition (TASCOG): The Tasmanian Study of Gait and Cognition is supported by project grants from the National Health and Medical Research Council of Australia (NHMRC; 403000,491109, and 606543) and a grant from the Wicking Dementia Education and Research Centre, Hobart. V.S. is supported by a cofunded NHMRC Career Development Fellowship (1061457) and a Heart Foundation Future Leader Fellowship (ID 100089).

Three-City Dijon Study: The Three-City Study is conducted under a partnership agreement among the Institut National de la Santé et de la Recherche Médicale (INSERM), the Victor Segalen–Bordeaux II University, and Sanofi-Aventis. The Fondation pour la Recherche Médicale funded the preparation and initiation of the study. The Three-City Study is also supported by the Caisse Nationale Maladie des Travailleurs Salariés, Direction Générale de la Santé, Mutuelle Générale de l'Education Nationale (MGEN), 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.” Christophe Tzourio and Stéphanie Debette are supported by a grant from the Fondation Leducq.

TOP: The study was supported by the Research Council of Norway (#213837, #223273, #229129), South-East Norway Health Authority (#2013-123) and KG Jebsen Foundation. The research leading to these results has received funding from the European Community's Seventh Framework Programme (FP7/2007-2013) under grant agreement no 602450 (IMAGEMEND).

UCLA_NL_BP: Data collection and genotyping was made possible with (NIH/NIMH) R01 MH090553 to R.A.O.th-East Norway Health Authority (#2013-123) and KG Jebsen Foundation.

UMCU: UMCU acknowledgment data: This work was supported by 917.46.370 (H.H.) and 908-02-123 (H.H.) from the Netherlands Organization for Health Research and Development ZonMW.

WHICAP: This study was supported by a grant from the NIH (5R01AG037212).

CHARGE consortium: See ref. 40 for the general cognitive function GWAS, and ref. 29 for the white matter lesion GWAS.

Early Growth Genetics (EGG) consortium: Data on head circumference, birth weight, and birth length have been contributed by the EGG Consortium and was downloaded from http://www.egg-consortium.org.

Genetic Investigation of ANthropometric Traits (GIANT) consortium: See ref. 13.

Genetics of Personality Consortium: See ref. 41 for the neuroticism GWAS and ref. 42 for the extraversion GWAS.

IGAP: We thank the International Genomics of Alzheimer's Project (IGAP) for providing summary results data for these analyses. The investigators within IGAP contributed to the design and implementation of IGAP and/or provided data but did not participate in analysis or writing of this report. IGAP was made possible by the generous participation of the control subjects, the patients, and their families. The i–Select chips was funded by the French National Foundation on Alzheimer's disease and related disorders. EADI was supported by the LABEX (laboratory of excellence program investment for the future) DISTALZ grant, Inserm, Institut Pasteur de Lille, Université de Lille 2 and the Lille University Hospital. GERAD was supported by the Medical Research Council (Grant no 503480), Alzheimer's Research UK (Grant no 503176), the Wellcome Trust (Grant no 082604/2/07/Z) and German Federal Ministry of Education and Research (BMBF): Competence Network Dementia (CND) grant no 01GI0102, 01GI0711, 01GI0420. CHARGE was partly supported by the NIH/NIA grant R01 AG033193 and the NIA AG081220 and AGES contract N01–AG–12100, the NHLBI grant R01 HL105756, the Icelandic Heart Association, and the Erasmus Medical Center and Erasmus University. ADGC was supported by the NIH/NIA grants: U01 AG032984, U24 AG021886, U01 AG016976, and the Alzheimer's Association grant ADGC–10–196728.

International Parkinson's Disease Genomics Consortium (IPDGC): See ref. 43.

Psychiatric Genomics Consortium: See ref. 44.

Social Science Genetic Association Consortium (SSGAC): See ref. 45 for the childhood cognitive function GWAS.

Author information

Author notes

    • Hieab H H Adams
    • , Derrek P Hibar
    • , Vincent Chouraki
    • , Jason L Stein
    • , Paul A Nyquist
    • , Miguel E Rentería
    • , Stella Trompet
    •  & Alejandro Arias-Vasquez

    These authors contributed equally to this work.

    • Margaret J Wright
    • , Lenore J Launer
    • , Gunter Schumann
    • , Myriam Fornage
    • , Barbara Franke
    • , Stéphanie Debette
    • , Sarah E Medland
    • , M Arfan Ikram
    •  & Paul M Thompson

    These authors jointly directed this work.

Affiliations

  1. Department of Epidemiology, Erasmus MC, Rotterdam, the Netherlands.

    • Hieab H H Adams
    • , Sven J Van der Lee
    • , Najaf Amin
    • , Albert Hofman
    • , M Kamran Ikram
    • , Henning Tiemeier
    • , Andre G Uitterlinden
    • , Cornelia M Van Duijn
    • , Meike W Vernooij
    •  & M Arfan Ikram
  2. Department of Radiology and Nuclear Medicine, Erasmus MC, Rotterdam, the Netherlands.

    • Hieab H H Adams
    • , Wiro J Niessen
    • , Aad Van der Lugt
    • , Meike W Vernooij
    • , Tonya White
    •  & M Arfan Ikram
  3. Imaging Genetics Center, USC Mark and Mary Stevens Neuroimaging and Informatics Institute, Keck School of Medicine of University of Southern California, Los Angeles, California, USA.

    • Derrek P Hibar
    • , Jason L Stein
    • , Neda Jahanshad
    • , Christopher R K Ching
    • , Christopher D Whelan
    •  & Paul M Thompson
  4. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, USA.

    • Vincent Chouraki
    • , Sudha Seshadri
    • , Alexa Beiser
    • , Claudia L Satizabal
    •  & Stéphanie Debette
  5. Lille University, Inserm, CHU Lille, Institut Pasteur de Lille, U1167 - RID-AGE - Risk factors and molecular determinants of aging-related diseases, Lille, France.

    • Vincent Chouraki
    •  & Philippe Amouyel
  6. Framingham Heart Study, Framingham, Massachusetts, USA.

    • Vincent Chouraki
    • , Sudha Seshadri
    • , Alexa Beiser
    • , Claudia L Satizabal
    •  & Anita L DeStefano
  7. Department of Genetics and UNC Neuroscience Center, University of North Carolina (UNC), Chapel Hill, North Carolina, USA.

    • Jason L Stein
  8. Department of Neurology, Department of Anesthesia/Critical Care Medicine, Department of Neurosurgery, Johns Hopkins University, Baltimore, Maryland, USA.

    • Paul A Nyquist
  9. QIMR Berghofer Medical Research Institute, Brisbane, Australia.

    • Miguel E Rentería
    • , Gabriel Cuellar-Partida
    • , Lachlan T Strike
    • , Narelle K Hansell
    • , Grant W Montgomery
    • , Nicholas G Martin
    •  & Sarah E Medland
  10. Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands.

    • Stella Trompet
    •  & J Wouter Jukema
  11. Department of Human Genetics, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Alejandro Arias-Vasquez
    • , Janita Bralten
    • , Martine Hoogman
    • , Marieke Klein
    • , Elena Shumskaya
    • , Marjolein M J Van Donkelaar
    • , Thomas Wolfers
    • , Hans van Bokhoven
    • , Han G Brunner
    •  & Barbara Franke
  12. Department of Psychiatry, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Alejandro Arias-Vasquez
    • , Nanda Rommelse
    •  & Barbara Franke
  13. Department of Cognitive Neuroscience, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Alejandro Arias-Vasquez
    • , Corina U Greven
    • , Jennifer S Richards
    • , Daan Van Rooij
    • , Jan K Buitelaar
    •  & Guillén Fernández
  14. Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, the Netherlands.

    • Alejandro Arias-Vasquez
    • , Janita Bralten
    • , Martine Hoogman
    • , Marieke Klein
    • , Andre Marquand
    • , Jennifer S Richards
    • , Nanda Rommelse
    • , Elena Shumskaya
    • , Marjolein M J Van Donkelaar
    • , Daan Van Rooij
    • , Thomas Wolfers
    • , Marcel P Zwiers
    • , Hans van Bokhoven
    • , Han G Brunner
    • , Jan K Buitelaar
    • , Guillén Fernández
    • , Simon E Fisher
    • , Clyde Francks
    •  & Barbara Franke
  15. MRC-SGDP Centre, Institute of Psychiatry, Psychology and Neuroscience, King's College London, London, UK.

    • Sylvane Desrivières
    • , Tianye Jia
    • , Christine Macare
    • , Bing Xu
    •  & Gunter Schumann
  16. Dr. John T. Macdonald Foundation Department of Human Genetics, University of Miami, Miller School of Medicine, Miami, Florida, USA.

    • Ashley H Beecham
    •  & Susan H Blanton
  17. John P. Hussman Institute for Human Genomics, University of Miami, Miller School of Medicine, Miami, Florida, USA.

    • Ashley H Beecham
    • , Susan H Blanton
    •  & Ralph L Sacco
  18. German Center for Neurodegenerative Diseases (DZNE) Rostock/Greifswald, Greifswald, Germany.

    • Katharina Wittfeld
    •  & Wolfgang Hoffmann
  19. Department of Psychiatry, University Medicine Greifswald, Greifswald, Germany.

    • Katharina Wittfeld
    • , Deborah Janowitz
    •  & Hans J Grabe
  20. Brain Center Rudolf Magnus, Department of Psychiatry, UMC Utrecht, Utrecht, the Netherlands.

    • Lucija Abramovic
    • , Marc M Bohlken
    • , Marco P Boks
    • , Rachel M Brouwer
    • , Wiepke Cahn
    • , Hilleke E Hulshoff Pol
    • , René S Kahn
    • , Roel A Ophoff
    •  & Neeltje E M Van Haren
  21. Department of Neurology and Neurosurgery, Montreal Neurological Institute, McGill University, Montreal, Canada.

    • Saud Alhusaini
  22. The Royal College of Surgeons in Ireland, Dublin 2, Ireland.

    • Saud Alhusaini
    • , Christopher D Whelan
    • , Gianpiero L Cavalleri
    •  & Norman Delanty
  23. Department of Integrative Medical Biology and Umeå center for Functional Brain Imaging, Umeå University, Umeå, Sweden.

    • Micael Andersson
    • , Sara Pudas
    •  & Lars Nyberg
  24. Department of Biomedical Engineering, Illinois Institute of Technology, Chicago, Illinois, USA.

    • Konstantinos Arfanakis
  25. Rush Alzheimer′s Disease Center, Rush University Medical Center, Chicago, Illinois, USA.

    • Konstantinos Arfanakis
    • , Jingyun Yang
    • , Neelum T Aggarwal
    • , David A Bennett
    •  & Debra A Fleischman
  26. Department of Diagnostic Radiology and Nuclear Medicine, Rush University Medical Center, Chicago, Illinois, USA.

    • Konstantinos Arfanakis
  27. Brain Research Imaging Centre, University of Edinburgh, Edinburgh, UK.

    • Benjamin S Aribisala
    • , Susana Muñoz Maniega
    • , Natalie A Royle
    • , Mark E Bastin
    • , Maria C Valdés Hernández
    •  & Joanna M Wardlaw
  28. Department of Computer Science, Lagos State University, Lagos, Nigeria.

    • Benjamin S Aribisala
  29. Scottish Imaging Network, A Platform for Scientific Excellence (SINAPSE) Collaboration, Department of Neuroimaging Sciences, University of Edinburgh, Edinburgh, UK.

    • Benjamin S Aribisala
    • , Susana Muñoz Maniega
    • , Natalie A Royle
    • , Mark E Bastin
    • , Maria C Valdés Hernández
    •  & Joanna M Wardlaw
  30. Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney, Australia.

    • Nicola J Armstrong
    • , Karen A Mather
    • , Amelia A Assareh
    • , Henry Brodaty
    • , Simone Reppermund
    • , Perminder S Sachdev
    • , Anbupalam Thalamuthu
    •  & Wei Wen
  31. Mathematics and Statistics, Murdoch University, Perth, Australia.

    • Nicola J Armstrong
  32. NORMENT - KG Jebsen Centre, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

    • Lavinia Athanasiu
    • , Nhat Trung Doan
    • , Unn K Haukvik
    • , Ingrid Agartz
    • , Ole A Andreassen
    • , Erik G Jönsson
    •  & Ingrid Melle
  33. NORMENT - KG Jebsen Centre, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.

    • Lavinia Athanasiu
    • , Lars T Westlye
    • , Ole A Andreassen
    • , Thomas Espeseth
    •  & Ingrid Melle
  34. Department of Medical Sciences, Molecular Medicine and Science for Life Laboratory, Uppsala University, Uppsala, Sweden.

    • Tomas Axelsson
  35. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA.

    • Alexa Beiser
  36. Hospital for Sick Children, University of Toronto, Toronto, Canada.

    • Manon Bernard
    • , Jean Shin
    •  & Zdenka Pausova
  37. Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, USA.

    • Joshua C Bis
  38. Generation R Study Group, Erasmus Medical Center, Rotterdam, the Netherlands.

    • Laura M E Blanken
    • , Ryan L Muetzel
    •  & Irene Pappa
  39. Department of Child and Adolescent Psychiatry/Psychology, Erasmus MC-Sophia Children′s Hospital, Rotterdam, the Netherlands.

    • Laura M E Blanken
    • , Ryan L Muetzel
    • , Henning Tiemeier
    •  & Tonya White
  40. Taub Institute for Research on Alzheimer's Disease and the Aging Brain, Columbia University Medical Center, New York, New York, USA.

    • Adam M Brickman
    • , Sandra Barral
    •  & Badri N Vardarajan
  41. G.H. Sergievsky Center, Columbia University Medical Center, New York, New York, USA.

    • Adam M Brickman
  42. Department of Neurology, Columbia University Medical Center, New York, New York, USA.

    • Adam M Brickman
  43. Pennington Biomedical Research Center, Baton Rouge, Louisiana, USA.

    • Owen Carmichael
  44. Cerebral Imaging Centre, Douglas Mental Health University Institute, Montreal, Canada.

    • M Mallar Chakravarty
  45. Department of Psychiatry and Biomedical Engineering, McGill University, Montreal, Canada.

    • M Mallar Chakravarty
  46. INSERM Unit U1219, University of Bordeaux, France.

    • Ganesh Chauhan
  47. Lieber Institute for Brain Development, Baltimore, Maryland, USA.

    • Qiang Chen
    • , Aaron L Goldman
    • , Venkata S Mattay
    • , Daniel R Weinberger
    •  & Stéphanie Debette
  48. Interdepartmental Neuroscience Graduate Program, UCLA School of Medicine, Los Angeles, California, USA.

    • Christopher R K Ching
  49. Biological Psychology, Neuroscience Campus Amsterdam, Vrije Universiteit University and Vrije Universiteit Medical Center, Amsterdam, the Netherlands.

    • Anouk Den Braber
    • , Dorret I Boomsma
    • , Eco J C De Geus
    • , Iryna O Fedko
    • , Jouke-Jan Hottenga
    •  & Dennis Van ′t Ent
  50. Division of Psychological and Social Medicine and Developmental Neurosciences, Faculty of Medicine, TU Dresden, Germany.

    • Stefan Ehrlich
  51. Department of Psychiatry, Massachusetts General Hospital, Boston, Masschusetts, USA.

    • Stefan Ehrlich
    • , Avram J Holmes
    • , Phil H Lee
    • , Randy L Buckner
    • , Randy L Gollub
    • , Joshua L Roffman
    •  & Jordan W Smoller
  52. Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Charlestown, North Carolina, USA.

    • Stefan Ehrlich
    • , Tian Ge
    •  & Randy L Gollub
  53. NSERM Unit 1000 ″Neuroimaging and Psychiatry″, University Paris Sud, University Paris Descartes, Paris, France.

    • Irina Filippi
    • , Hervé Lemaître
    •  & Jean-Luc Martinot
  54. Maison de Solenn, Adolescent Psychopathology and Medicine Department, APHP Hospital Cochin, Paris, France.

    • Irina Filippi
    • , Hervé Lemaître
    •  & Jean-Luc Martinot
  55. Psychiatric and Neurodevelopmental Genetics Unit, Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Tian Ge
    • , Phil H Lee
    •  & Jordan W Smoller
  56. Harvard Medical School, Boston, Massachusetts, USA.

    • Tian Ge
    • , Phil H Lee
    • , Philip L De Jager
    • , Randy L Gollub
    • , Robert C Green
    •  & Jordan W Smoller
  57. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Boston, Massachusetts, USA.

    • Tian Ge
    • , Phil H Lee
    •  & Jordan W Smoller
  58. NORMENT - KG Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Norway.

    • Sudheer Giddaluru
    • , Srdjan Djurovic
    • , Stephanie Le Hellard
    •  & Vidar M Steen
  59. Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway.

    • Sudheer Giddaluru
    • , Anita L DeStefano
    • , Stephanie Le Hellard
    •  & Vidar M Steen
  60. Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Rebecca F Gottesman
    •  & Venkata S Mattay
  61. Karakter Child and Adolescent Psychiatry University Center, Nijmegen, the Netherlands.

    • Corina U Greven
    • , Jennifer S Richards
    • , Nanda Rommelse
    •  & Jan K Buitelaar
  62. King′s College London, Medical Research Council Social, Genetic and Developmental Psychiatry Centre, Institute of Psychology, Psychiatry and Neurosciene, London, UK.

    • Corina U Greven
  63. Central Institute of Mental Health, Medical Faculty Mannheim, University Heidelberg, Mannheim, Germany.

    • Oliver Grimm
    • , Andreas Meyer-Lindenberg
    •  & Marcella Rietschel
  64. Center of Biostatistics and Bioinformatics, University of Mississippi Medical Center, Jackson, Mississippi, USA.

    • Michael E Griswold
  65. Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, the Netherlands.

    • Tulio Guadalupe
    • , Simon E Fisher
    •  & Clyde Francks
  66. International Max Planck Research School for Language Sciences, Nijmegen, the Netherlands.

    • Tulio Guadalupe
  67. Department of Child and Adolescent Psychiatry, Faculty of Medicine of the TU Dresden, Dresden, Germany.

    • Johanna Hass
    •  & Esther Walton
  68. Department of Research and Development, Diakonhjemmet Hospital, Oslo, Norway.

    • Unn K Haukvik
    •  & Ingrid Agartz
  69. Department of Pharmacology, National University of Singapore, Singapore.

    • Saima Hilal
    • , Christopher Chen
    •  & M Kamran Ikram
  70. Memory Aging and Cognition Centre (MACC), National University Health System, Singapore.

    • Saima Hilal
    • , Christopher Chen
    •  & M Kamran Ikram
  71. Department of Neurology, Clinical Division of Neurogeriatrics, Medical University Graz, Austria, Graz, Austria.

    • Edith Hofer
    • , Lukas Pirpamer
    •  & Reinhold Schmidt
  72. Institute of Medical Informatics, Statistics and Documentation, Medical University Graz, Austria, Graz, Austria.

    • Edith Hofer
  73. Max Planck Institute of Psychiatry, Department of Translational Research in Psychiatry, Munich, Germany.

    • David Hoehn
    • , Nazanin Mirza-Schreiber
    • , Benno Pütz
    • , Philipp G Sämann
    • , Michael Czisch
    • , Florian Holsboer
    •  & Bertram Müller-Myhsok
  74. Department of Psychology, Yale University, New Haven, Connecticut, USA.

    • Avram J Holmes
  75. UCL Institute of Neurology, London, United Kingdom and Epilepsy Society, Bucks, UK.

    • Dalia Kasperaviciute
    • , Mar Matarin
    •  & Sanjay M Sisodiya
  76. Department of Medicine, Imperial College London, London, UK.

    • Dalia Kasperaviciute
  77. Center for Neuroimaging, Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Sungeun Kim
    • , Kwangsik Nho
    • , Shannon L Risacher
    • , Li Shen
    •  & Andrew J Saykin
  78. Center for Computational Biology and Bioinformatics, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Sungeun Kim
    • , Kwangsik Nho
    • , Li Shen
    •  & Tatiana M Foroud
  79. Indiana Alzheimer Disease Center, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Sungeun Kim
    • , Kwangsik Nho
    • , Shannon L Risacher
    • , Li Shen
    •  & Andrew J Saykin
  80. Section for Experimental Psychopathology and Neuroimaging, Department of General Psychiatry, Heidelberg University, Heidelberg, Germany.

    • Bernd Kraemer
    •  & Oliver Gruber
  81. Lurie Center for Autism, Massachusetts General Hospital, Harvard Medical School, Lexington, Massachusetts, USA.

    • Phil H Lee
  82. Singapore Eye Research Institute, Singapore National Eye Centre, Singapore.

    • Jiemin Liao
    • , Ching-Yu Cheng
    • , M Kamran Ikram
    •  & Tien Y Wong
  83. Centre for Cognitive Ageing and Cognitive Epidemiology, Psychology, University of Edinburgh, Edinburgh, UK.

    • David C M Liewald
    • , Lorna M Lopez
    • , Michelle Luciano
    • , Susana Muñoz Maniega
    • , Natalie A Royle
    • , Mark E Bastin
    • , Andrew M McIntosh
    • , Maria C Valdés Hernández
    • , Joanna M Wardlaw
    •  & Ian J Deary
  84. Donders Centre for Cognitive Neuroimaging, Radboud University, Nijmegen, The Netherlands.

    • Andre Marquand
    • , Elena Shumskaya
    •  & Marcel P Zwiers
  85. Reta Lila Weston Institute and Department of Molecular Neuroscience, UCL Institute of Neurology, London, UK.

    • Mar Matarin
    • , Adaikalavan Ramasamy
    • , Daniah Trabzuni
    • , J Raphael Gibbs
    • , Sebastian Guelfi
    • , John Hardy
    • , Dena G Hernandez
    •  & Mina Ryten
  86. Department of Biomedicine, Aarhus University, Aarhus, Denmark.

    • Manuel Mattheisen
  87. The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Aarhus and Copenhagen, Denmark.

    • Manuel Mattheisen
  88. Center for integrated Sequencing, iSEQ, Aarhus University, Aarhus, Denmark.

    • Manuel Mattheisen
  89. UMR5296 University of Bordeaux, CNRS, CEA, Bordeaux, France.

    • Bernard Mazoyer
  90. Department of Psychiatry, Yale University, New Haven, Connecticut, USA.

    • David R McKay
    • , Emma Sprooten
    • , Anderson M Winkler
    •  & David C Glahn
  91. Olin Neuropsychiatric Research Center, Hartford, Connecticut, USA.

    • David R McKay
    • , Emma Sprooten
    •  & David C Glahn
  92. Menzies Institute for Medical Research, University of Tasmania, Hobart, Tasmania, Australia.

    • Rebekah McWhirter
  93. Department of Psychiatry, EMGO Institute for Health and Care Research and Neuroscience Campus Amsterdam, VU University Medical Center/GGZ inGeest, Amsterdam, The Netherlands.

    • Yuri Milaneschi
  94. Experimental Therapeutics and Pathophysiology Branch, National Institute of Mental Health Intramural Research Program, National Institutes of Health, Bethesda, Maryland, USA.

    • Allison C Nugent
    • , Girma Woldehawariat
    • , Dara M Cannon
    • , Wayne C Drevets
    • , Xinmin Liu
    •  & Francis J McMahon
  95. Center for Neurobehavioral Genetics, University of California, Los Angeles, California, USA.

    • Loes M Olde Loohuis
    •  & Roel A Ophoff
  96. Department of Clinical Neuropsychology, VU University Amsterdam, Amsterdam, the Netherlands.

    • Jaap Oosterlaan
  97. Division of Psychiatry, Royal Edinburgh Hospital, University of Edinburgh, Edinburgh, UK.

    • Martina Papmeyer
    • , Emma Sprooten
    • , Stephen M Lawrie
    • , Andrew M McIntosh
    •  & Jessika E Sussmann
  98. Division of Systems Neuroscience of Psychopathology, Translational Research Center, University Hospital of Psychiatry, University of Bern, Switzerland.

    • Martina Papmeyer
  99. School of Pedagogical and Educational Sciences, Erasmus University Rotterdam, Rotterdam, the Netherlands.

    • Irene Pappa
  100. Rush Institute for Healthy Aging, Rush University Medical Center, Chicago, Illinois, USA.

    • Kumar B Rajan
    • , Neelum T Aggarwal
    •  & Denis A Evans
  101. Department of Medical and Molecular Genetics, King's College London, London, UK.

    • Adaikalavan Ramasamy
    • , Mina Ryten
    •  & Michael E Weale
  102. The Jenner Institute Laboratories, University of Oxford, Oxford, UK.

    • Adaikalavan Ramasamy
  103. Department of Medicine and Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria-IDIVAL, Santander, Spain.

    • Roberto Roiz-Santiañez
    •  & Benedicto Crespo-Facorro
  104. CIBERSAM (Centro Investigación Biomédica en Red Salud Mental), Santander, Spain.

    • Roberto Roiz-Santiañez
    • , Diana Tordesillas-Gutierrez
    •  & Benedicto Crespo-Facorro
  105. Psychosis Research Group, Department of Psychiatry and Trinity Translational Medicine Institute, Trinity College Dublin.

    • Emma J Rose
    •  & Aiden Corvin
  106. Centre for Clinical Brain Sciences, University of Edinburgh, Edinburgh, UK.

    • Natalie A Royle
    • , Mark E Bastin
    • , Maria C Valdés Hernández
    •  & Joanna M Wardlaw
  107. Department of Neurology, University of Miami, Miller School of Medicine, Miami, Florida, USA.

    • Tatjana Rundek
    • , Ralph L Sacco
    •  & Clinton B Wright
  108. Department of Epidemiology and Public Health Sciences, University of Miami, Miller School of Medicine, Miami, Florida, USA.

    • Tatjana Rundek
    • , Ralph L Sacco
    •  & Clinton B Wright
  109. Orygen, The National Centre of Excellence in Youth Mental Health, Melbourne, VIC, Australia.

    • Lianne Schmaal
  110. Centre for Youth Mental Health, The University of Melbourne, Melbourne, VIC, Australia.

    • Lianne Schmaal
  111. Department of Psychiatry, Neuroscience Campus Amsterdam, VU University Medical Center, Amsterdam, the Netherlands.

    • Lianne Schmaal
    • , Brenda W J H Penninx
    •  & Dick J Veltman
  112. Multimodal Imaging Laboratory, Department of Neurosciences, University of California, San Diego, USA.

    • Andrew J Schork
  113. Department of Cognitive Sciences, University of California, San Diego, USA.

    • Andrew J Schork
  114. Icelandic Heart Association, Kopavogur, Iceland.

    • Albert V Smith
    • , Vilmundur Gudnason
    •  & Sigurdur Sigurdsson
  115. Faculty of Medicine, University of Iceland, Reykjavik, Iceland.

    • Albert V Smith
    •  & Vilmundur Gudnason
  116. Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Emma Sprooten
  117. Queensland Brain Institute, University of Queensland, Brisbane, Australia.

    • Lachlan T Strike
    • , Narelle K Hansell
    •  & Margaret J Wright
  118. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Alexander Teumer
    • , Wolfgang Hoffmann
    •  & Henry Völzke
  119. School of Computing Engineering and Mathematics, Western Sydney University, Parramatta, Australia.

    • Russell Thomson
  120. Neuroimaging Unit,Technological Facilities. Valdecilla Biomedical Research Institute IDIVAL, Santander, Cantabria, Spain.

    • Diana Tordesillas-Gutierrez
  121. Institut Pasteur, Paris, France.

    • Roberto Toro
  122. Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia.

    • Daniah Trabzuni
  123. GeneSTAR Research Center, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Dhananjay Vaidya
    • , Lisa R Yanek
    •  & Diane M Becker
  124. Department of Radiology, Leiden University Medical Center, Leiden, the Netherlands.

    • Jeroen Van der Grond
  125. Department of Psychiatry, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Dennis Van der Meer
    • , Daan Van Rooij
    • , Catharina A Hartman
    •  & Pieter J Hoekstra
  126. Brain Center Rudolf Magnus, Human Neurogenetics Unit, UMC Utrecht, Utrecht, the Netherlands.

    • Kristel R Van Eijk
  127. Department of Psychiatry and Human Behavior, University of California-Irvine, Irvine, California, USA.

    • Theo G M Van Erp
    •  & Steven G Potkin
  128. NORMENT - KG Jebsen Centre, Department of Psychology, University of Oslo, Oslo, Norway.

    • Lars T Westlye
    •  & Thomas Espeseth
  129. Department of Medicine, University of Mississippi Medical Center, Jackson, Mississippi, USA.

    • Beverly G Windham
    •  & Thomas H Mosley
  130. FMRIB Centre, University of Oxford, Oxford, UK.

    • Anderson M Winkler
    • , Mark Jenkinson
    • , Erik G Jönsson
    •  & Thomas E Nichols
  131. University of Wuerzburg, Department of Psychiatry, Psychosomatics and Psychotherapy, Wuerzburg, Germany.

    • Christiane Wolf
  132. Department of Neurological Sciences, Rush University Medical Center, Chicago, Illinois, USA.

    • Jingyun Yang
    • , Neelum T Aggarwal
    •  & David A Bennett
  133. Biospective Inc, Montreal, Quebec, Canada, Montréal, Québec, Canada.

    • Alex Zijdenbos
  134. Department of Clinical Neuroscience, Centre for Psychiatric Research, Karolinska Institutet, Stockholm, Sweden.

    • Ingrid Agartz
  135. South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine Brownsville/Edinburg/San Antonio, Texas, USA.

    • Laura Almasy
    • , John Blangero
    • , Joanne E Curran
    • , Ravi Duggirala
    • , Thomas D Dyer
    •  & Harald H H Göring
  136. Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA.

    • Laura Almasy
  137. Department of Biomedical and Health Informatics, The Children′s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA.

    • Laura Almasy
  138. National Ageing Research Institute, Royal Melbourne Hospital, Melbourne, Australia.

    • David Ames
  139. Academic Unit for Psychiatry of Old Age, University of Melbourne, Melbourne, Australia.

    • David Ames
  140. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, USA.

    • Sampath Arepalli
    • , Mark R Cookson
    • , Allissa Dillman
    • , J Raphael Gibbs
    • , Dena G Hernandez
    • , Michael A Nalls
    • , Andrew Singleton
    •  & Bryan J Traynor
  141. Departments of Psychiatry, Neurology, and Psychology, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • James T Becker
  142. Dementia Collaborative Research Centre - Assessment and Better Care, UNSW, Sydney, Australia.

    • Henry Brodaty
  143. Department of Clinical Genetics, Maastricht University Medical Center, Maastricht, the Netherlands.

    • Han G Brunner
  144. Department of Psychology, Center for Brain Science, Harvard University, Cambridge, Massachusetts, USA.

    • Randy L Buckner
  145. Department of Evolution and Genetics, Dagestan State University, Makhachkala, Dagestan, Russia.

    • Kazima B Bulayeva
    •  & Tien Y Wong
  146. The Mind Research Network and LBERI, Albuquerque, New Mexico, USA.

    • Vince D Calhoun
  147. Department of ECE, University of New Mexico, Albuquerque, New Mexico, USA.

    • Vince D Calhoun
  148. Centre for Neuroimaging and Cognitive Genomics (NICOG), Clinical Neuroimaging Laboratory, NCBES Galway Neuroscience Centre, College of Medicine Nursing and Health Sciences, National University of Ireland Galway, Galway, Ireland.

    • Dara M Cannon
    • , Colm McDonald
    •  & Thomas W Mühleisen
  149. Academic Medicine Research Institute, Duke-NUS Graduate Medical School, Singapore.

    • Ching-Yu Cheng
    •  & M Kamran Ikram
  150. Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore.

    • Ching-Yu Cheng
    •  & Tien Y Wong
  151. Division of Medical Genetics, Department of Biomedicine, University of Basel, Basel, Switzerland.

    • Sven Cichon
  152. Institute of Human Genetics, University of Bonn, Bonn, Germany.

    • Sven Cichon
    •  & Markus M Nöthen
  153. Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany.

    • Sven Cichon
    •  & Thomas W Mühleisen
  154. Center for Multimodal Imaging and Genetics, University of California, San Diego, California, USA.

    • Anders M Dale
  155. Department of Neurosciences, University of California, San Diego, California, USA.

    • Anders M Dale
  156. Department of Radiology, University of California, San Diego, California, USA.

    • Anders M Dale
  157. Department of Psychiatry, University of California, San Diego, California, USA.

    • Anders M Dale
  158. Department of Cognitive Science, University of California, San Diego, California, USA.

    • Anders M Dale
  159. Avera Institute for Human Genetics, Sioux Falls, South Dakota, USA.

    • Gareth E Davies
  160. Program in Translational NeuroPsychiatric Genomics, Departments of Neurology and Psychiatry, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Gareth E Davies
  161. Harvard Medical School, Boston, Massachusetts, USA.

    • Gareth E Davies
  162. Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, USA.

    • Gareth E Davies
    •  & Philip L De Jager
  163. Broad Institute, Cambridge, Massachusetts, USA.

    • Philip L De Jager
  164. Faculty of Health and Institute of Health and Biomedical Innovation, Queensland University of Technology (QUT), Brisbane, Australia.

    • Greig I de Zubicaray
  165. Neurology Division, Beaumont Hospital, Dublin, 9, Ireland.

    • Norman Delanty
  166. Department of Neurology, Hopital Erasme, Universite Libre de Bruxelles, Brussels, Belgium.

    • Chantal Depondt
    •  & Massimo Pandolfo
  167. Department of Medical Genetics, Oslo University Hospital, Oslo, Norway.

    • Srdjan Djurovic
  168. Cognitive Genetics and Cognitive Therapy Group, Neuroimaging, Cognition and Genomics Centre (NICOG) and NCBES Galway Neuroscience Centre, School of Psychology and Discipline of Biochemistry, National University of Ireland Galway, Galway, Ireland.

    • Gary Donohoe
    •  & Derek W Morris
  169. Neuropsychiatric Genetics Research Group, Department of Psychiatry and Trinity College Institute of Psychiatry, Trinity College Dublin, Dublin 8, Ireland.

    • Gary Donohoe
    •  & Derek W Morris
  170. Janssen Research and Development, LLC, Titusville, New Jersey, USA.

    • Wayne C Drevets
  171. Department of Psychiatry and Psychotherapy, Charité Universitätsmedizin Berlin, CCM, Berlin, Germany.

    • Susanne Erk
    • , Andreas Heinz
    • , Sebastian Mohnke
    • , Nina Romanczuk-Seiferth
    •  & Henrik Walter
  172. Intramural Research Program of the National Institute on Aging, Baltimore, Maryland, USA.

    • Luigi Ferrucci
  173. Department of Neurological Sciences, Rush University Medical Center, Chicago, Illinois, USA.

    • Debra A Fleischman
  174. Department of Behavioral Sciences, Rush University Medical Center, Chicago, Illinois, USA.

    • Debra A Fleischman
  175. Robertson Center for Biostatistics, University of Glasgow, Glasgow, UK.

    • Ian Ford
  176. Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Tatiana M Foroud
    •  & Andrew J Saykin
  177. University of Texas Health Science Center, San Antonio, Texas, USA.

    • Peter T Fox
    •  & Rene L Olvera
  178. Division of Cerebral Integration, National Institute for Physiological Sciences, Aichi, Japan.

    • Masaki Fukunaga
  179. Division of Genetics, Department of Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Robert C Green
  180. Department of Psychiatry, Osaka University Graduate School of Medicine, Osaka, Japan.

    • Ryota Hashimoto
    •  & Kazutaka Ohi
  181. Molecular Research Center for Children′s Mental Development, United Graduate School of Child Development, Osaka University, Osaka, Japan.

    • Ryota Hashimoto
  182. Institute of Diagnostic Radiology and Neuroradiology, University Medicine Greifswald, Greifswald, Germany.

    • Katrin Hegenscheid
    •  & Norbert Hosten
  183. German Center for Neurodegenerative Diseases (DZNE), Tübingen, Germany.

    • Dena G Hernandez
  184. Department of Psychology, VU University Amsterdam, Amsterdam, the Netherlands.

    • Dirk J Heslenfeld
  185. Department of Psychiatry, University of Iowa, Iowa City, Iowa, USA.

    • Beng-Choon Ho
  186. HMNC Brain Health, Munich, Germany.

    • Florian Holsboer
  187. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, Germany.

    • Georg Homuth
  188. Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Japan.

    • Masashi Ikeda
  189. Department of Radiology, Mayo Clinic, Rochester, Minnesota, USA.

    • Clifford R Jack Jr
  190. NICHD Brain and Tissue Bank for Developmental Disorders, University of Maryland Medical School, Baltimore, Maryland, USA.

    • Robert Johnson
    •  & H Ronald Zielke
  191. School of Psychology, University of Sussex, Brighton, UK.

    • Ryota Kanai
  192. Institute of Cognitive Neuroscience, University College London, London, UK.

    • Ryota Kanai
  193. Department of Neuroinformatics, Araya Brain Imaging, Tokyo, Japan.

    • Ryota Kanai
  194. Medical University of Lodz, Lodz, Poland.

    • Iwona Kloszewska
  195. Department of Neurology, Mayo Clinic, Rochester, Minnesota, USA.

    • David S Knopman
  196. Maryland Psychiatric Research Center, Department of Psychiatry, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Peter Kochunov
  197. Neuroscience Research Australia, Sydney, Australia.

    • John B Kwok
    •  & Peter R Schofield
  198. School of Medical Sciences, UNSW, Sydney, Australia.

    • John B Kwok
    •  & Peter R Schofield
  199. Columbia University Medical Center, New York, New York, USA.

    • Xinmin Liu
  200. Laboratory of Genetics, National Institute on Aging, National Institutes of Health, Baltimore, Maryland, USA.

    • Dan L Longo
  201. Department of Neurology, University of Washington, Seattle, Washington, USA.

    • W T Longstreth Jr
  202. Department of Epidemiology, University of Washington, Seattle, Washington, USA.

    • W T Longstreth Jr
  203. Department of Neurology, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • Oscar L Lopez
  204. Department of Psychiatry, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • Oscar L Lopez
  205. Department of Psychiatry, University of Oxford, Oxford, UK.

    • Simon Lovestone
  206. NIHR Dementia Biomedical Research Unit, King's College London, London, UK.

    • Simon Lovestone
  207. Imaging of Dementia and Aging (IDeA) Laboratory, Department of Neurology and Center for Neuroscience, University of California at Davis, Sacramento, California, USA.

    • Oliver Martinez
    •  & Charles DeCarli
  208. Department of Radiology, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Venkata S Mattay
  209. Centre for Advanced Imaging, University of Queensland, Brisbane, Australia.

    • Katie L McMahon
    •  & Margaret J Wright
  210. Section of Gerontology and Geriatrics, Department of Medicine, University of Perugia, Perugia, Italy.

    • Patrizia Mecocci
  211. Munich Cluster for Systems Neurology (SyNergy), Munich, Germany.

    • Bertram Müller-Myhsok
  212. University of Liverpool, Institute of Translational Medicine, Liverpool, UK.

    • Bertram Müller-Myhsok
  213. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Matthias Nauck
  214. German Center for Cardiovascular Research (DZHK e.V.), partner site Greifswald, Germany.

    • Matthias Nauck
  215. Department of Statistics and Warwick Manufacturing Group, University of Warwick, Coventry, UK.

    • Thomas E Nichols
  216. Department of Medical Informatics Erasmus MC, Rotterdam, the Netherlands.

    • Wiro J Niessen
  217. Faculty of Applied Sciences, Delft University of Technology, Delft, the Netherlands.

    • Wiro J Niessen
  218. Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany.

    • Markus M Nöthen
  219. Rotman Research Institute, University of Toronto, Toronto, Canada.

    • Tomas Paus
  220. Departments of Psychology and Psychiatry, University of Toronto, Canada.

    • Tomas Paus
  221. Child Mind Institute, New York, New York, USA.

    • Tomas Paus
  222. Department of Physiology, University of Toronto, Toronto, Canada.

    • Zdenka Pausova
  223. Department of Nutritional Sciences, University of Toronto, Toronto, Canada.

    • Zdenka Pausova
  224. Department of Radiology, University of Calgary, Calgary, Canada.

    • G Bruce Pike
  225. Department of Clinical Neuroscience, University of Calgary, Calgary, Canada.

    • G Bruce Pike
  226. Department of Epidemiology, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  227. Department of Medicine, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  228. Department of Health Services, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  229. Group Health Research Institute, Group Health, Seattle, Washington, USA.

    • Bruce M Psaty
  230. Department of Developmental Disability Neuropsychiatry, School of Psychiatry, UNSW Medicine, Australia.

    • Simone Reppermund
  231. Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute and Pediatrics at Harbor-UCLA Medical Center, Torrance, California, USA.

    • Jerome I Rotter
  232. Evelyn F. McKnight Brain Institute, University of Miami, Miller School of Medicine, Miami, Florida, USA.

    • Ralph L Sacco
    •  & Clinton B Wright
  233. Neuropsychiatric Institute, Prince of Wales Hospital, Sydney, Australia.

    • Perminder S Sachdev
  234. Department of Neuroimaging, Institute of Psychiatry, King's College London, London, UK.

    • Andy Simmons
  235. Biomedical Research Centre for Mental Health, King's College London, London, UK.

    • Andy Simmons
  236. Biomedical Research Unit for Dementia, King's College London, London, UK.

    • Andy Simmons
  237. MRC Edinburgh Brain Bank, University of Edinburgh, Academic Department of Neuropathology, Centre for Clinical Brain Sciences, Edinburgh, UK.

    • Colin Smith
  238. Institute of Clinical Medicine, Neurology, University of Eastern Finland, Kuopio, Finland.

    • Hilkka Soininen
  239. Neurocentre Neurology, Kuopio University Hospital, Finland.

    • Hilkka Soininen
  240. Department of Medicine, Peninsula Health and Monash University, Melbourne, Australia.

    • Velandai Srikanth
  241. Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, UK.

    • David J Stott
  242. Laboratory of Neuro Imaging, Institute for Neuroimaging and Informatics, Keck School of Medicine of the University of Southern California, Los Angeles, California, USA.

    • Arthur W Toga
  243. Brain Resource Center, Johns Hopkins University, Baltimore, Maryland, USA.

    • Juan Troncoso
  244. Georgia State University, Atlanta, Georgia, USA.

    • Jessica A Turner
  245. Institute for Neurodegenerative Disorders, UMR 5293, CEA, CNRS, Université de Bordeaux, France.

    • Christophe Tzourio
  246. Department of Internal Medicine, Erasmus MC, Rotterdam, the Netherlands.

    • Andre G Uitterlinden
  247. Genentech Inc., South San Francisco, California, USA.

    • Marcel Van der Brug
  248. Department of Psychiatry and Leiden Institute for Brain and Cognition, Leiden University Medical Center, Leiden, The Netherlands.

    • Nic J A Van der Wee
  249. Neuroimaging Centre, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Marie-Jose Van Tol
  250. Department of Psychiatry, Carver College of Medicine, University of Iowa, Iowa City, Iowa, USA.

    • Thomas H Wassink
  251. Departments of Psychiatry, Neurology, Neuroscience and the Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Daniel R Weinberger
  252. Center for Imaging of Neurodegenerative Disease, San Francisco VA Medical Center, University of California, San Francisco, California, USA.

    • Michael W Weiner
  253. Department of Neurobiology, Care Sciences and Society, Karolinska Institutet, Stockholm, Sweden.

    • Eric Westman
  254. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, USA.

    • Alan B Zonderman
  255. Institute of Molecular Biology and Biochemistry, Medical University Graz, Graz, Austria.

    • Helena Schmidt
  256. Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands.

    • Anton J M De Craen
  257. Intramural Research Program, NIA, NIH, Bethesda, Maryland, USA.

    • Lenore J Launer
  258. Institute of Molecular Medicine and Human Genetics Center, University of Texas Health Science Center at Houston, Houston, Texas, USA.

    • Myriam Fornage
  259. Department of Neurology, Bordeaux University Hospital, Bordeaux, France.

    • Stéphanie Debette
  260. Department of Neurology, Erasmus MC, Rotterdam, the Netherlands.

    • M Arfan Ikram

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Contributions

Conceived of the study and drafted the manuscript: H.H.H.A., D.P.H., V.C., J.L.S., M.E.R., S.T., A.A., P.N, V.G., G.S., M.F., B.F., S.D., S.E.M., M.A.I., P.M.T. Performed statistical analyses: H.H.H.A., D.P.H., V.C., J.L.S., M.E.R., S.T., A.A., Sy.D., A.H.B., N.J., K.W., Lu.A., N.A., M.A., B.S.A., N.J.A., La.A., A.B., M.B., J.C.B., L.M.E.B., S.H.B., M.M.B., Ja.B., O.C., M.M.C., Ga.C., Q.C., C.R.K.C., G.C., Nh.D., St.E., Ti.G., Su.G., A.L.G., C.U.G., Ol.G., M.E.G., T.G., Jo.H., U.K.H., S.H., E.H., M.H., D.J., T.J., N.K., D.K., S.K., M.K., B.K., P.H.L., J.L., D.C.M.L., L.M.L., M.L., Ch.M., Su.M., An.M., Ma.M., M.M., Be.M., D.R.M., R.M., Y.M., R.L.M., K.N., L.M.O., J.O., Ma.P., I.P., L.P., S.P., B.P., K.B.R., A.R., J.S.R., S.L.R., R.R., Na.R., N.A.R., T.R., C.L.S., Li.S., An.J.S., L.S., J.S., A.V.S., E.S., L.T.S., Al.T., Ro.T., Di.T., R.T., D.T., Dh.V., J.V., S.J.V., D.vdM., M.M.J.V., K.R.V., D.vR., Es.W., L.T.W., A.M.W., G.W., C.W., Th.W., L.R.Y., J.Y., M.P.Z., A.M.D., I.O.F., B.M., T.E.N., J.A.T., B.X., Sa.A., A.M.B., A.dB., A.J.H., A.C.N., P.G.S., C.D.W., S.M.B., R.M.B., G.D., J.G., O.G., R.K., C.M., M.A.N., D.V., B.N.V., T.W., E.J.R. Acquired data: P.N., Su.S., K.A.A., T.A., M.P.B., Ir.F., R.F.G., D.H., K.A.M., Em.S., B.G.W., A.Z., I.A., N.T.A., L.A., D.A., P.A., O.A.A., S.A., A.A.A., M.E.B., D.M.B., J.T.B., D.A.B., J.B., H.v.a.B., D.I.B., H.B., H.G.B., R.L.B., J.K.B., K.B.B., W.C., V.D.C., D.M.C., G.L.C., C-Y.C., C.C., S.C., M.R.C., A.C., B.C., J.E.C., M.C., G.E.D., E.J.C.D., P.L.D., G.I.D., N.D., Ch.D., A.DeS., A.D., Sr.D., W.C.D., R.D., T.D.D., S.E., T.E., D.A.E., G.F., L.F., S.E.F., D.A.F., I.F., T.M.F., P.T.F., C.F., Ma.F., D.C.G., R.L.G., H.H.H.G., H.J.G., R.C.G., S.G., N.K.H., J.H., C.A.H., R.H., K.H., An.H., S.L.e.H., D.G.H., D.J.H., B.H., P.J.H., W.H., A.H., F.H., G.H., N.H., J-J.H., H.E.H., M.I., M.K.I., C.R.J., R.J., E.G.J., J.J., R.S.K., I.K., D.S.K., P.K., J.B.K., L.J.L., S.M.L., H.L., X.L., D.L.L., W.L., O.L.L., S.L., O.M., J.M., V.S.M., A.M.M., F.J.M., K.L.M., P.M., I.M., A.M., S.M., G.W.M., D.W.M., T.H.M., T.W.M., M.N., W.J.N., M.M.N., L.N., K.O., R.L.O., R.A.O., M.P., T.P., Z.P., B.W.J.P., G.P., S.G.P., B.M.P., S.R., Ma.R., J.L.R., N.R., J.I.R., M.R., R.L.S., P.S.S., A.J.S., He.S., P.R.S., S.S., A.S., S.M.S., C.S., J.W.S., H.S., V.S., V.M.S., D.J.S., J.E.S., A.T., H.T., A.W.T., B.T., J.T., C.T., A.G.U., M.C.V., M.vdB., A.V., N.J.A.V., C.M.V., N.E.M.V., M.V., D.J.V., M.W.V., H.V., H.W., J.M.W., T.H.W., M.E.W., D.R.W., M.W.W., W.W., E.W., T.Y.W., C.B.W., R.H.Z., A.B.Z., I.J.D., C.D., R.S., N.G.M., A.J.M.D., M.J.W., V.G., G.S., M.F., B.F., S.D., S.E.M., M.A.I., P.M.T., A.Sim., Sa.A., A.M.B., A.dB., A.J.H., A.C.N., P.G.S., C.D.W., S.M.B., R.M.B., G.D., J.G., O.G., R.K., C.M., M.A.N., D.V., B.N.V., T.W., E.J.R. All authors critically reviewed the manuscript for important intellectual content.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to M Arfan Ikram or Paul M Thompson.

Integrated supplementary information

Supplementary information

PDF files

  1. 1.

    Supplementary Text and Figures

    Supplementary Figures 1–3

  2. 2.

    Supplementary Methods Checklist

Excel files

  1. 1.

    Supplementary Table 1

    Information on sampling and demographics of study populations.

  2. 2.

    Supplementary Table 2

    Information on genotyping and quality control.

  3. 3.

    Supplementary Table 3

    Information on image acquisition and processing.

  4. 4.

    Supplementary Table 4

    Functional annotation of genome-wide significant variants associated with intracranial volume and all variants in LD (> 0.8).

  5. 5.

    Supplementary Table 5

    Phenotypic correlation between intracranial volume and height.

  6. 6.

    Supplementary Table 6

    Genome-wide significant variants associated with intracranial volume with and without adjustment for height.

  7. 7.

    Supplementary Table 7

    The association between a polygenic score constructed from the 697 variants identified for height by the GIANT consortium and intracranial volume, with and without adjustment for height.

  8. 8.

    Supplementary Table 8

    Genome-wide significant variants associated with height and their association with intracranial volume.

  9. 9.

    Supplementary Table 9

    Novel genome-wide significant variants after meta-analysis of intracranial volume and child head circumference.

  10. 10.

    Supplementary Table 10

    Pathway analysis of intracranial volume using the KGG software.

  11. 11.

    Supplementary Table 11

    Pathway analysis of intracranial volume using the MAGENTA software.

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DOI

https://doi.org/10.1038/nn.4398