Abstract
Copy number variants (CNVs) may increase the risk for neurodevelopmental conditions. The neurobiological mechanisms that link these high-risk genetic variants to clinical phenotypes are largely unknown. An important question is whether brain abnormalities in individuals who carry CNVs are associated with their degree of penetrance.
Methods
We investigated whether increased CNV penetrance for schizophrenia and other developmental disorders was associated with variations in cortical and subcortical morphology. We pooled T1-weighted brain magnetic resonance imaging and genetic data from 22 cohorts from the ENIGMA (Enhancing Neuro Imaging Genetics through Meta Analysis)-CNV consortium. In the main analyses, we included 9268 individuals (aged 7–90 years, 54% female), from which we identified 398 carriers of 36 neurodevelopmental CNVs at 20 distinct loci. A secondary analysis was performed including additional neuroimaging data from the ENIGMA-22q consortium, including 274 carriers of the 22q11.2 deletion and 291 noncarriers. CNV penetrance was estimated through penetrance scores that were previously generated from large cohorts of patients and controls. These scores represent the probability risk of developing either schizophrenia or other developmental disorders (including developmental delay, autism spectrum disorder, and congenital malformations).
Results
For both schizophrenia and developmental disorders, increased penetrance scores were associated with lower surface area in the cerebral cortex and lower intracranial volume. For both conditions, associations between CNV-penetrance scores and cortical surface area were strongest in regions of the occipital lobes, specifically in the cuneus and lingual gyrus.
Conclusions
Our findings link global and regional cortical morphometric features with CNV penetrance, providing new insights into neurobiological mechanisms of genetic risk for schizophrenia and other developmental disorders.
| Original language | English |
|---|---|
| Pages (from-to) | 1093-1106 |
| Number of pages | 14 |
| Journal | Biological Psychiatry: Cognitive Neuroscience and Neuroimaging |
| Volume | 10 |
| Issue number | 10 |
| Early online date | 23 May 2025 |
| DOIs | |
| Publication status | Published - Oct 2025 |
Funding
METH-CT: DJS is supported by the South African Medical Research Council (SAMRC). FS is supported by a PDRF (Presidential Doctoral Research Fellowship) from the Brain Behavior Unit, Department of Psychiatry and Mental Health, University of Cape Town. SD is supported by the Biomedical Research and Innovation Platform and SAMRC. NCNG: NCNG (Norwegian Cognitive NeuroGenetics sample) was supported by the Bergen Research Foundation , the University of Bergen , the Research Council of Norway (FUGE) (Enhance Research in Functional Genomics) (Grant Nos. 151904 and 183327), Psykisk Helse (Grant No. 175345), RCN (Grant Nos. 154313/V50 [to Ivar Reinvang] and 177458/V50 [to TE]), Helse S\u00F8r\u00D8st RHF (Grant No. 2012086 [to TE]), and the Dr. Einar Martens Fund. ECHO-DEFINE: The ECHO (Experiences of Children with Copy Number Variants) study acknowledges funding from the Wellcome Trust (Institutional Strategic Support Fund) (to MBMvdB) and Clinical Research Training Fellowship (Grant No. 102003/Z/13/Z [to JLD]), the Waterloo Foundation (Grant No. WF 918- 1234 [to MBMvdB]), the Baily Thomas Charitable Fund (Grant No. 2315/1 [to MBMvdB]), National Institute of Mental Health (NIMH) (Grant No. 5UO1MH101724 ) and NIMH (Grant No. U01MH119738 [to MBMvdB]), the IMAGINE-ID and IMAGINE-2 studies (funded by Medical Research Council [MRC]) (Grant Nos. MR/N022572/1 and MR/T033045/1 [to MBMvdB]) and an MRC Centre Grant (Grant No. MR/P005748/1 [to MJO]). The DEFINE (Defining Endophenotypes From Integrative Neuroscience) study was supported by a Wellcome Trust Strategic Award (Grant No. 100202/Z/12/Z [to MJO]). NTR: The NTR cohort was supported by the Netherlands Organization for Scientific Research (NWO) and The Netherlands Organization for Health Research and Development (ZonMW) (Grant Nos. 904-61-090, 985-10-002, 912-10-020, 904-61-193, 480-04-004, 463-06-001, 451-04-034, 400-05-717, Addiction-31160008, 016-115-035, 481-08-011, 400-07-080, 056-32-010, Middelgroot-911-09-032, OCW_NWO Gravity program \u2013024.001.003, NWO-Groot 480-15-001/674), Center for Medical Systems Biology (CMSB, NWO Genomics), NBIC/BioAssist/RK(2008.024), Biobanking and Biomolecular Resources Research Infrastructure (BBMRI) (Grant Nos. NL, 184.021.007 and 184.033.111), X-Omics 184-034-019; Spinozapremie (Grant No. NWO- 56-464-14192), KNAW Academy Professor Award (PAH/6635) and University Research Fellow grant (to DIB); Amsterdam Public Health Research Institute (former EMGO+), Neuroscience Amsterdam research institute (former NCA); the European Community\u2019s Fifth and Seventh Framework Program (Grant Nos. FP5- LIFE QUALITY-CT-2002-2006, FP7-HEALTH-F4-2007-2013, Grant No. 01254: GenomEUtwin, Grant No. 01413: ENGAGE and Grant No. 602768:ACTION); the European Research Council (Grant Nos. ERC Starting 284\u2006167, ERC Consolidator 771057, ERC Advanced 230374), Rutgers University Cell and DNA Repository (NIMH) (Grant No. U24 MH068457-06), NIH (Grant Nos. R01D0042157-01A1, R01MH58799-03, MH081802, DA018673, R01DK092127-04), Grand Opportunity (Grant Nos. 1RC2 MH089951 and 1RC2 MH089995); and the Avera Institute for Human Genetics , Sioux Falls, South Dakota. Part of the genotyping and analyses were funded by the Genetic Association Information Network of the Foundation for the NIH. Computing was supported by NWO (Grant No. 2018/EW/00408559), BiG Grid, the Dutch e-Science Grid, and SURFSARA. IMAGEN: The European Union\u2013funded FP6 Integrated Project IMAGEN (Reinforcement-related behavior in normal brain function and psychopathology) (Grant No. LSHM-CT- 2007-037286), the Horizon 2020 funded ERC Advanced Grant STRATIFY (Brain network based stratification of reinforcement-related disorders) (Grant No. 695313), Human Brain Project (Grant Nos. HBP SGA 2, 785907, and HBP SGA 3, 945539), the MRC Grant c-VEDA (Consortium on Vulnerability to Externalizing Disorders and Addictions) (Grant No. MR/N000390/1), NIH (Grant No. R01DA049238), A decentralized macro and micro gene-by-environment interaction analysis of substance use behavior and its brain biomarkers), the NIHR Biomedical Research Centre at South London and Maudsley NHS Foundation Trust and King\u2019s College London, the Bundesministerium f\u00FCr Bildung und Forschung (Grant Nos. 01GS08152 ; 01EV0711; Forschungsnetz AERIAL 01EE1406A, 01EE1406B; Forschungsnetz IMAC- Mind 01GL1745B), the Deutsche Forschungsgemeinschaft (DFG Grant Nos. SM 80/7-2 , SFB 940 , TRR 265 , NE 1383/14-1 ), ImagenPathways \u201CUnderstanding the Interplay between Cultural, Biological and Subjective Factors in Drug Use Pathways\u201D is a collaborative project supported by the European Research Area Network on Illicit Drugs, the Medical Research Foundation and MRC (Grant Nos. MR/R00465X/1 and MR/S020306/1), the NIH-funded ENIGMA (Grant Nos. 5U54EB020403-05 and 1R56AG058854-01), National Natural Science Foundation of China (NSFC) (Grant No. 82\u2006150\u2006710\u2006554), and European Union\u2013funded project environMENTAL (Grant No. 101057429). Additional support was provided by grants from the ANR (Grant Nos. ANR-12-SAMA-0004, AAPG2019\u2014GeBra), the Eranet Neuron (Grant Nos. AF12-NEUR0008-01\u2014WM2NA; and ANR-18-NEUR00002-01\u2014ADORe), the Fondation de France (Grant No. 00081242 ), the Fondation pour la Recherche M\u00E9dicale (Grant No. DPA20140629802), the Mission Interminist\u00E9rielle de Lutte-contre-les-Drogues-et-les-Conduites-Addictives, the Assistance Publique-H\u00F4pitaux de Paris and Institut National de la Sant\u00E9 et de la Recherche M\u00E9dicale (INSERM) (interface grant), Paris Sud University IDEX 2012, the Fondation de l\u2019Avenir (Grant No. AP-RM-17-013), the F\u00E9d\u00E9ration pour la Recherche sur le Cerveau; the NIH, Science Foundation Ireland (Grant No. 16/ERCD/3797 ), the Science Foundation Ireland (Grant No. 16/ERCD/3797), the NIH (Axon, Testosterone and Mental Health during Adolescence) (Grant No. R01MH085772-01A1), and the NIH Consortium (Grant No. U54 EB020403), supported by a cross-NIH alliance that funds Big Data to Knowledge Centres of Excellence. The INSERM, the Strasbourg University, and SATT CONECTUS, provided sponsorship (principal investigator: J-LM). TP is supported by NIH (Grant No. R01MH085772). Gilles Berstchy is acknowledged for his support. Stephane Lehericy and the radiographer staff at Centre de NeuroImagerie de Recherche de l'\u2019Institut du Cerveau ( http://www.cenir.org/mri.html?lang=en ) are acknowledged for their support in acquisition of MRI datasets. SHIP-2/SHIP-TREND: SHIP is part of the Community Medicine Research net of the University of Greifswald , Germany, which is funded by the Federal Ministry of Education and Research (Grant Nos. 01ZZ9603 , 01ZZ0103 , and 01ZZ0403 ), the Ministry of Cultural Affairs as well as the Social Ministry of the Federal State of Mecklenburg-West Pomerania, and the network \u201CGreifswald Approach to Individualized Medicine (GANI_MED)\u201D funded by the Federal Ministry of Education and Research (Grant No. 03IS2061A ). Genome-wide data have been supported by the Federal Ministry of Education and Research (Grant No. 03ZIK012 ) and a joint grant from Siemens Healthineers and the Federal State of Mecklenburg-West Pomerania. The University of Greifswald is a member of the Cach\u00E9 Campus program of the InterSystems GmbH. This study has received funding from the following institutions: Federal Ministry of Education and Research, the Ministry of Cultural Affairs, as well as the Social Ministry of the Federal State of Mecklenburg-West Pomerania. MRI examinations were supported by Siemens Healthineers, Siemens Healthcare GmbH. AT has been funded by DFG (Grant No. 542489987). The SHIP authors thank Holger Prokisch and Thomas Meitinger (Helmholtz Zentrum M\u00FCnchen) for the genotyping of the SHIP-TREND cohort. 16p11.2 Consortium: This work was supported by Compute Canada , the Brain Canada Multi investigator research initiative, Canada First Research Excellence Fund, the Institute of Data Valorization, and Healthy Brain Healthy Lives. SJ is a recipient of a Canada Research Chair in neurodevelopmental disorders and a chair from the Jeanne et Jean Louis Levesque Foundation. This work was supported by the Canadian Institutes of Health Research (CIHR) (Grant Nos. 400528 and 438531 ) and Innovative Medicines Initiative 2 Joint Undertaking (Grant No. 777394 ) for AIMS-2-TRIALS. This Joint Undertaking receives support from the EU\u2019s Horizon 2020 program, EFPIA , Autism Speaks , Autistica , and SFARI . KK was supported by The Institute of Data Valorization Postdoctoral Fellowship program , through the Canada First Research Excellence Fund. GAP: PD\u2019s research is supported by the MRC and the Wellcome Trust. Funding was received from the Psychiatry Research Trust, the Wellcome Trust, the MRC, and the National Institute for Health Research (NIHR) Biomedical Research Centre at the South London and Maudsley NHS Trust. The funding sources were not involved in carrying out or interpreting the study. OATS1/OATS2 and Sydney MAS: The OATS (Older Australian Twins Study) has been funded by a National Health & Medical Research Council (NHMRC) and Australian Research Council (ARC) Strategic Award Grant of the Ageing Well, Ageing Productively Program (Grant No. 401162); NHMRC Project (seed) Grants (Grant Nos. 1024224 and 1\u2006025\u2006243); NHMRC Project Grants (Grant Nos. 1045325 and 1085606); and NHMRC Program Grants (Grant Nos. 568969 and 1093083). This research was facilitated through access to Twins Research Australia, which is a national resource supported by a Centre of Research Excellence Grant (Grant No. 1079102) from the NHMRC. The Sydney Memory and Ageing Study has been funded by 3 NHMRC Program Grants (Grant Nos. ID350833, ID568969, and APP1093083). Core funding was received from the NSW Government Department of Health and the Australian government Department of Health and Aged Care; NHMRC Leadership Fellowship (Grant No. GNT2009771 ); NHMRC (Grant No. GNT1045325). DNA samples were extracted by Genetic Repositories Australia, an Enabling Facility, which was supported by an NHMRC Grant (Grant No. 401184). MRI scans were processed with the support of NHMRC Project Grants (Grant Nos. 510175 and 1025243) and an ARC Discovery Project Grant (Grant No. DP0774213) and John Holden Family Foundation. We thank the participants and their informants for their time and generosity in contributing to this research and acknowledge the contributions of the respective research teams. HUNT: The HUNT (Tr\u00F8ndelag Health Study) is a collaboration between HUNT Research Centre (Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology), North Tr\u00F8ndelag County Council, Central Norway Health Authority, and the Norwegian Institute of Public Health. HUNT-MRI was funded by the Liaison Committee between the Central Norway Regional Health Authority and the Norwegian University of Science and Technology and the Norwegian National Advisory Unit for functional MRI. HUBIN (Human Brain Informatics): This research is supported by Swedish Research Council (Grant Nos. K2007-62X-15077-04-1 , K2008-62P-20597-01-3 , K2010-62X-15078-07-2 , K2012-61X-15078-09-3 , K2015-62X-15077-12-3 , and 2017-00949 ), the regional agreement on medical training and clinical research between Stockholm County Council and the Karolinska Institutet, and the Research Council of Norway (Grant No. 274\u2006359). PAFIP: This study was supported by grants from Carlos III Health Institute (Grant Nos. IPI17/00402 , PI17/01056, PI14/00639, and PI14/00918) cofunded by the European Union through FEDER funds and Fundaci\u00F3n Instituto de Investigaci\u00F3n Marqu\u00E9s de Valdecilla (Grant Nos. NCT0235832 and NCT02534363). No pharmaceutical company has financially supported the study. JV-B acknowledges IDIVAL Grants (Grant Nos. INT/A21/10 and INT/A20/04). Haavik: This research is supported by Stiftelsen KG Jebsen (Grant No. SKGJ MED-02), the Research Council of Norway, and Helse Vest RHF. ENIGMA: The ENIGMA Working Group acknowledges the NIH Big Data to Knowledge award for foundational support and consortium development (Grant No. U54 EB020403 [to PMT]). For a complete list of ENIGMA-related grant support, please see here: http://enigma.ini.usc.edu/about-2/funding/ . PMT and SIT are also supported in part by ENIGMA NIH (Grant Nos. R01MH116147, R01NS107513, R01AG058854, and R01MH129742). IES is supported by the Research Council of Norway (Grant No. 223273 ), South-Eastern Norway Regional Health Authority (Grant No. 2020060 ), European Union\u2019s Horizon2020 Research and Innovation Programme (CoMorMent project) (Grant No. 847776 ), and Kristian Gerhard Jebsen Stiftelsen (Grant No. SKGJ-MED-021). South-Eastern Norway Regional Health Authority (Grant No. 2020060 ) also supports RBo. This work was partially performed on Services for Sensitive Data (TSD), University of Oslo, Norway, with resources provided by UNINETT Sigma2, the National Infrastructure for High Performance Computing and Data Storage in Norway. OAA is supported by Research Council of Norway (Grant No. 223273) and KG Jebsen Stiftelsen (Grant No. SKGJ-MED-021). NJ is supported by NIH (Grant No. R01MH117601). DvdM is supported by Research Council of Norway (Grant No. 276082). IA has received lecturer honorarium from Lundbeck. TB served in an advisory or consultancy role for eye level, InfectoPharm, Lundbeck, Medice, Neurim Pharmaceuticals, Oberberg GmbH, Roche, and Takeda. TB received conference support or speaker\u2019s fee by Janssen, Medice, and Takeda. TB received royalties from Hogrefe, Kohlhammer, CIP Medien, and Oxford University Press; the current work is unrelated to these relationships. PD has received speaker\u2019s fees from Lundbeck and Janssen. TRM has received honoraria for speaking and chairing from Lundbeck, Janssen, Astellas, and Viatris and received honoraria to participate in advisory boards organized by Angelini Pharmaceuticals. TRM is an employee of and shareholder in Pasithea Therapeutics. OAA is a consultant for HealthLytix and received speaker\u2019s honorarium from Lundbeck and Sunovion. MJO is in receipt of research grants from Takeda Pharmaceuticals and Akrivia Health outside of the scope of the current work. MBMvdB is in receipt of a research grant from Takeda Pharmaceuticals outside of the scope of the current work. BAG is a paid consultant for Natera, Inc. JHaa has received lecture honoraria as part of continuing medical education programs sponsored by Shire, Takeda, Medice, and Biocodex. PSS was an expert panel member for Biogen Australia and Roche Australia in 2020 and 2021. HJG has received speaker\u2019s honoraria from Servier, Neurapharm, Indorsia, and Janssen-Cilag. JASV has served as a consultant for NoBias Therapeutics Inc. and has received speaker fees for giving a lecture in the Henry Stewart Talks series. LP has received speaking fees from InfectoPharm, Medice, and Takeda and royalties from Hogrefe, Kohlhammer, and Schattauer. The current work is unrelated to these relationships. All other authors report no biomedical financial interests or potential conflicts of interest. MCIC (Mind Clinical Imaging Consortium): This research is funded by NIH study P20GM103472 . Stroke (TOP_T3): The European Research Council under the European Union\u2019s Horizon 2020 research and Innovation program (Grant No. ERC 802998). The Research Council of Norway (Grant Nos. 298646 and 300767), and the South-Eastern Norway Regional Health Authority (Grant Nos. 2013054 , 2015044 , 2015073 , 2018076 , 2019101 , and 2020060 ). QTIM: The QTIM study was supported by grants from the U.S. National Institute of Child Health and Human Development (Grant No. R01HD050735 ) and the Australian NHMRC (Grant Nos. 486682 , 1009064 ). Genotyping was supported by NHMRC (Grant No. 389875). AFM is funded by an ARC Future Fellowship (Grant No. FT200100837). SEM is supported by NHMRC Investigator Grant (Grant No. APP1172917). ENIGMA-22q: This research is supported by NIMH (Grant Nos. R01MH085953, R21MH116473, and 9U01MH119736-02). ASB is supported by the Dalglish Family Chair in 22q11.2 Deletion Syndrome, CIHR (Grant Nos. MOP-79518, MOP-89066, MOP-97800, and MOP-111238) and NIMH (Grant No. U01MH101723\u201301[3/5]). EWCC acknowledges support from the CIHR (Grant No. MOP-74631) and Ontario Mental Health Foundation for the current work. DRR is supported by NIMH (Grant No. R01 119185). JES is supported by the Stanford Maternal and Child Health Research Institute Uytengsu-Hamilton 22q11 Neuropsychiatry Research Awards Program (Grant No. UH22QEXTFY21). DMM-M acknowledges support from NIMH (Grant No. U01MH119737). JASV acknowledges funding support from NIMH (Grant Nos. 1U01MH119741-01 and 3U01MH119741-02S1). TvA acknowledges financial support from NIMH (Grant No. 5U01 MH119740). Dublin: This research is funded by European Research Council (Grant No. 677467 ). GOBS: The GOBS (Genetics of Brain Structure and Function study) data collection was supported in part by NIH (Grant Nos. R01MH078143 and R01MH083824). COBRE (Centers of Biomedical Research Excellence): This research is funded by National Science Foundation (Grant No. 2112455 ) and National Institutes of Health (NIH) (Grant No. R01MH118695 ).
| Funders | Funder number |
|---|---|
| South African Medical Research Council | |
| NCNG | |
| IMAGINE-ID | |
| Netherlands Organization for Scientific Research | |
| Paris Sud University | |
| Siemens Healthineers | |
| Institut de Valorisation des Données | |
| National Institute for Health Research | |
| John Holden Family Foundation | |
| Amsterdam Public Health Research Institute | |
| European Federation of Pharmaceutical Industries and Associations | |
| EU’s Horizon 2020 program | |
| King’s College London | |
| Assistance Publique-Hôpitaux de Paris | |
| Central Norway Regional Health Authority | |
| Biomedical Research and Innovation Platform | |
| SURFsara | |
| University of Cape Town | |
| PDRF | |
| Medical Research Council | |
| Ministry of Cultural Affairs | |
| Biomedical Research Centre at the South London and Maudsley NHS Trust | |
| Neuroscience Amsterdam research institute | |
| BiG Grid | |
| Karolinska Institutet | |
| FEDER | |
| Deutsche Forschungsgemeinschaft | |
| National Health & Medical Research Council | |
| Dr. Einar Martens Fund | |
| Helse Vest RHF | |
| Institut national de la santé et de la recherche médicale | |
| Fédération pour la Recherche sur le Cerveau | |
| Canada First Research Excellence Fund | |
| Autism Speaks | |
| Stockholm County Council | |
| National Natural Science Foundation of China | |
| Netherlands Organization for Health Research and Development | |
| University of Bergen | |
| Compute Canada | |
| Simons Foundation Autism Research Initiative | |
| NWO | |
| Norwegian University of Science and Technology | |
| Psychiatry Research Trust | |
| NIHR Biomedical Research Centre at South London and Maudsley NHS Foundation Trust | |
| European Union | |
| Norway Regional Health Authority | |
| Mission Interministérielle de Lutte-contre-les-Drogues-et-les-Conduites-Addictives | |
| Takeda Pharmaceuticals | |
| Horizon 2020 | |
| NSFC | 82 150 710 554, 101057429 |
| NIH | 1RC2 MH089995, 1RC2 MH089951, DA018673, R01DA049238, R01MH078143, MH081802, R01D0042157-01A1, R01MH083824, R01MH58799-03, R01DK092127-04, P20GM103472 |
| Kristian Gerhard Jebsen Stiftelsen | SKGJ-MED-021 |
| Baily Thomas Charitable Fund | 2315/1 |
| Research Council of Norway | 223273, 300767, 298646, 274 359 |
| EraNet Neuron | ANR-18-NEUR00002-01—ADORe, AF12-NEUR0008-01—WM2NA |
| U.S. National Institute of Child Health and Human Development | R01HD050735 |
| FP6 | LSHM-CT- 2007-037286 |
| Bundesministerium für Bildung und Forschung | 01GS08152, 01EV0711, AERIAL 01EE1406A, 01EE1406B |
| Wellcome Trust | 102003/Z/13/Z |
| MRC | MR/T033045/1, MR/N022572/1, MR/N000390/1, MR/P005748/1, 100202/Z/12/Z |
| ZonMw | 016-115-035, 480-04-004, 463-06-001, 481-08-011, 451-04-034, 024.001.003, 400-07-080, 912-10-020, 904-61-090, 400-05-717, 904-61-193, 056-32-010, 985-10-002 |
| Stiftelsen KG Jebsen | SKGJ MED-02 |
| KG Jebsen Stiftelsen | R01MH117601, 276082 |
| NWO-Groot | 480-15-001/674 |
| Ontario Mental Health Foundation | R01 119185 |
| Stanford Maternal and Child Health Research Institute Uytengsu-Hamilton 22q11 Neuropsychiatry Research | U01MH119737, 5U01 MH119740, 3U01MH119741-02S1, 1U01MH119741-01, UH22QEXTFY21 |
| ENGAGE | 602768 |
| ANR | AAPG2019—GeBra, ANR-12-SAMA-0004 |
| Innovative Medicines Initiative | AIMS-2-TRIALS, 777394 |
| Biobanking and Biomolecular Resources Research Infrastructure | 184.033.111, NWO- 56-464-14192, X-Omics 184-034-019, 184.021.007 |
| ERC | 695313, 785907, 230374, 945539 |
| National Institute of Mental Health | U01MH119738, 5UO1MH101724 |
| Helse SørØst RHF | 2012086 |
| RCN | 154313/V50, 177458/V50 |
| Psykisk Helse | 175345 |
| Canadian Institutes of Health Research | 400528, MOP-97800, MOP-74631, MOP-111238, U01MH101723–01[3/5, MOP-79518, 438531, MOP-89066 |
| European Community’s Fifth and Seventh Framework Program | FP5- LIFE QUALITY-CT-2002-2006 |
| Waterloo Foundation | WF 918- 1234 |
| European Union’s Horizon2020 research and innovation programme | 847776 |
| IDIVAL | INT/A20/04, INT/A21/10 |
| FUGE | 183327, 151904 |
| NIH Consortium | R01MH085772, U54 EB020403 |
| Carlos III Health Institute | IPI17/00402, PI17/01056, PI14/00639, PI14/00918 |
| Avera Institute for Human Genetics | 2018/EW/00408559 |
| South-Eastern Norway Regional Health Authority | 2019101, 2015073, 2015044, 2013054, 2018076, 2020060 |
| IMAC | 01GL1745B |
| NBIC/BioAssist/RK | 2008.024 |
| Fundación Instituto de Investigación Marqués de Valdecilla | NCT0235832, NCT02534363 |
| European Research Council | 677467 |
| Australian Research Council | 1024224, 568969, ID568969, 510175, ID350833, GNT1045325, 1093083, GNT2009771, 1085606, 1079102, APP1093083, 401162, 401184, DP0774213, 1045325, 1 025 243 |
| DFG | SM 80/7-2, NE 1383/14-1, SFB 940, 542489987, TRR 265 |
| NIMH | R21MH116473, R01MH085953, U24 MH068457-06, 9U01MH119736-02 |
| Federal State of Mecklenburg-West Pomerania | 03ZIK012, 03IS2061A |
| European Union’s Horizon 2020 research and innovation program | ERC 802998 |
| National Science Foundation | 2112455 |
| FP7-HEALTH-F4-2007-2013 | 01254, 01413 |
| Science Foundation Ireland | 16/ERCD/3797, R01MH085772-01A1 |
| Medical Research Foundation | MR/R00465X/1, MR/S020306/1 |
| Fondation pour la Recherche Médicale | DPA20140629802 |
| KNAW | PAH/6635 |
| National Institutes of Health | R01MH118695 |
| Fondation de France | 00081242 |
| Federal Ministry of Education and Research | 01ZZ0403, 01ZZ0103, 01ZZ9603 |
| NHMRC | 486682, 389875, 1009064 |
| Swedish Research Council | K2007-62X-15077-04-1, K2015-62X-15077-12-3, K2010-62X-15078-07-2, K2012-61X-15078-09-3, 2017-00949, K2008-62P-20597-01-3 |
| ENIGMA NIH | R01MH129742, R01MH116147, R01AG058854, R01NS107513 |
| Fondation de l’Avenir | AP-RM-17-013 |
| ARC | APP1172917, FT200100837 |
| NIH-funded | 5U54EB020403-05, 1R56AG058854-01 |