{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T14:34:23Z","timestamp":1792593263695,"version":"4.5.0"},"reference-count":98,"publisher":"Springer Science and Business Media LLC","issue":"12","license":[{"start":{"date-parts":[[2025,11,11]],"date-time":"2025-11-11T00:00:00Z","timestamp":1762819200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2025,11,11]],"date-time":"2025-11-11T00:00:00Z","timestamp":1762819200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Neurosci"],"published-print":{"date-parts":[[2025,12]]},"DOI":"10.1038\/s41593-025-02090-2","type":"journal-article","created":{"date-parts":[[2025,11,11]],"date-time":"2025-11-11T10:02:56Z","timestamp":1762855376000},"page":"2439-2450","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":26,"title":["A genome-wide analysis of the shared genetic risk architecture of complex neurological and psychiatric disorders"],"prefix":"10.1038","volume":"28","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3761-5215","authenticated-orcid":false,"given":"Olav B.","family":"Smeland","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6602-2622","authenticated-orcid":false,"given":"Gleda","family":"Kutrolli","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8179-2788","authenticated-orcid":false,"given":"Shahram","family":"Bahrami","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5490-2390","authenticated-orcid":false,"given":"Vera","family":"Fominykh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8369-0152","authenticated-orcid":false,"given":"Nadine","family":"Parker","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6286-5614","authenticated-orcid":false,"given":"Julian","family":"Fuhrer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1666-4701","authenticated-orcid":false,"given":"Guy F. L.","family":"Hindley","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5967-6601","authenticated-orcid":false,"given":"Linn","family":"R\u00f8devand","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9408-3107","authenticated-orcid":false,"given":"Piotr","family":"Jaholkowski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3351-701X","authenticated-orcid":false,"given":"Markos","family":"Tesfaye","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1930-1167","authenticated-orcid":false,"given":"Pravesh","family":"Parekh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9991-1610","authenticated-orcid":false,"given":"Torbj\u00f8rn","family":"Elvs\u00e5shagen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7852-9244","authenticated-orcid":false,"given":"Andrew D.","family":"Grotzinger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6442-1179","authenticated-orcid":false,"given":"Nils Eiel","family":"Steen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dennis","family":"van der Meer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kevin S.","family":"O\u2019Connell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8140-8061","authenticated-orcid":false,"given":"Srdjan","family":"Djurovic","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6126-2966","authenticated-orcid":false,"given":"Anders M.","family":"Dale","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexey A.","family":"Shadrin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6427-2625","authenticated-orcid":false,"given":"Oleksandr","family":"Frei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4461-3568","authenticated-orcid":false,"given":"Ole A.","family":"Andreassen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,11,11]]},"reference":[{"key":"2090_CR1","doi-asserted-by":"publisher","first-page":"e0116820","DOI":"10.1371\/journal.pone.0116820","volume":"10","author":"HA Whiteford","year":"2015","unstructured":"Whiteford, H. A., Ferrari, A. J., Degenhardt, L., Feigin, V. & Vos, T. The global burden of mental, neurological and substance use disorders: an analysis from the Global Burden of Disease Study 2010. PLoS ONE 10, e0116820 (2015).","journal-title":"PLoS ONE"},{"key":"2090_CR2","unstructured":"WHO. International Classification of Diseases for Mortality and Morbidity Statistics, 11th revision (WHO, 2018)."},{"key":"2090_CR3","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1176\/ajp.155.4.457","volume":"155","author":"ER Kandel","year":"1998","unstructured":"Kandel, E. R. A new intellectual framework for psychiatry. Am. J. Psychiatry 155, 457\u2013469 (1998).","journal-title":"Am. J. Psychiatry"},{"key":"2090_CR4","doi-asserted-by":"publisher","first-page":"554","DOI":"10.1001\/jama.2020.0062","volume":"324","author":"MS Keshavan","year":"2020","unstructured":"Keshavan, M. S., Price, B. H. & Martin, J. B. The convergence of neurology and psychiatry: the importance of cross-disciplinary education. JAMA 324, 554\u2013555 (2020).","journal-title":"JAMA"},{"key":"2090_CR5","doi-asserted-by":"publisher","first-page":"2221","DOI":"10.1001\/jama.294.17.2221","volume":"294","author":"TR Insel","year":"2005","unstructured":"Insel, T. R. & Quirion, R. Psychiatry as a clinical neuroscience discipline. JAMA 294, 2221\u20132224 (2005).","journal-title":"JAMA"},{"key":"2090_CR6","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1038\/s44220-023-00086-0","volume":"1","author":"A Ibanez","year":"2023","unstructured":"Ibanez, A. & Zimmer, E. R. Time to synergize mental health with brain health. Nat. Ment. Health 1, 441\u2013443 (2023).","journal-title":"Nat. Ment. Health"},{"key":"2090_CR7","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1176\/appi.neuropsych.21120312","volume":"35","author":"JC Brown","year":"2023","unstructured":"Brown, J. C. et al. Time for brain medicine. J. Neuropsychiatry Clin. Neurosci. 35, 333\u2013340 (2023).","journal-title":"J. Neuropsychiatry Clin. Neurosci."},{"key":"2090_CR8","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1038\/nrd3628","volume":"11","author":"MJ Millan","year":"2012","unstructured":"Millan, M. J. et al. Cognitive dysfunction in psychiatric disorders: characteristics, causes and the quest for improved therapy. Nat. Rev. Drug Discov. 11, 141\u2013168 (2012).","journal-title":"Nat. Rev. Drug Discov."},{"key":"2090_CR9","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1038\/s41582-021-00597-3","volume":"18","author":"Z Ismail","year":"2022","unstructured":"Ismail, Z. et al. Psychosis in Alzheimer disease\u2014mechanisms, genetics and therapeutic opportunities. Nat. Rev. Neurol. 18, 131\u2013144 (2022).","journal-title":"Nat. Rev. Neurol."},{"key":"2090_CR10","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1038\/nrneurol.2016.200","volume":"13","author":"DH Ffytche","year":"2017","unstructured":"Ffytche, D. H. et al. The psychosis spectrum in Parkinson disease. Nat. Rev. Neurol. 13, 81\u201395 (2017).","journal-title":"Nat. Rev. Neurol."},{"key":"2090_CR11","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1111\/j.1600-0404.2004.00324.x","volume":"110","author":"A Gaitatzis","year":"2004","unstructured":"Gaitatzis, A., Trimble, M. R. & Sander, J. W. The psychiatric comorbidity of epilepsy. Acta Neurol. Scand. 110, 207\u2013220 (2004).","journal-title":"Acta Neurol. Scand."},{"key":"2090_CR12","doi-asserted-by":"publisher","first-page":"956","DOI":"10.1093\/schbul\/sbx089","volume":"43","author":"V Peralta","year":"2017","unstructured":"Peralta, V. & Cuesta, M. J. Motor abnormalities: from neurodevelopmental to neurodegenerative through \u2018functional\u2019 (neuro)psychiatric disorders. Schizophr. Bull. 43, 956\u2013971 (2017).","journal-title":"Schizophr. Bull."},{"key":"2090_CR13","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1136\/jnnp-2015-312233","volume":"87","author":"MT Minen","year":"2016","unstructured":"Minen, M. T. et al. Migraine and its psychiatric comorbidities. J. Neurol. Neurosurg. Psychiatry 87, 741\u2013749 (2016).","journal-title":"J. Neurol. Neurosurg. Psychiatry"},{"key":"2090_CR14","doi-asserted-by":"publisher","first-page":"l4485","DOI":"10.1136\/bmj.l4485","volume":"366","author":"V Shanker","year":"2019","unstructured":"Shanker, V. Essential tremor: diagnosis and management. BMJ 366, l4485 (2019).","journal-title":"BMJ"},{"key":"2090_CR15","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1001\/jamapsychiatry.2021.4377","volume":"79","author":"LS Richmond-Rakerd","year":"2022","unstructured":"Richmond-Rakerd, L. S., D\u2019Souza, S., Milne, B. J., Caspi, A. & Moffitt, T. E. Longitudinal associations of mental disorders with dementia: 30-year analysis of 1.7 million New Zealand citizens. JAMA Psychiatry 79, 333\u2013340 (2022).","journal-title":"JAMA Psychiatry"},{"key":"2090_CR16","doi-asserted-by":"publisher","DOI":"10.1038\/s41398-020-0705-1","volume":"10","author":"PM Thompson","year":"2020","unstructured":"Thompson, P. M. et al. ENIGMA and global neuroscience: a decade of large-scale studies of the brain in health and disease across more than 40 countries. Transl. Psychiatry 10, 100 (2020).","journal-title":"Transl. Psychiatry"},{"key":"2090_CR17","doi-asserted-by":"publisher","first-page":"1362","DOI":"10.1038\/s41380-021-01410-9","volume":"27","author":"E Leung","year":"2022","unstructured":"Leung, E. et al. Alterations in brain synaptic proteins and mRNAs in mood disorders: a systematic review and meta-analysis of postmortem brain studies. Mol. Psychiatry 27, 1362\u20131372 (2022).","journal-title":"Mol. Psychiatry"},{"key":"2090_CR18","doi-asserted-by":"publisher","first-page":"176","DOI":"10.1016\/j.tins.2022.12.004","volume":"46","author":"H Hampel","year":"2023","unstructured":"Hampel, H. et al. The foundation and architecture of precision medicine in neurology and psychiatry. Trends Neurosci. 46, 176\u2013198 (2023).","journal-title":"Trends Neurosci."},{"key":"2090_CR19","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1038\/ng.3285","volume":"47","author":"TJ Polderman","year":"2015","unstructured":"Polderman, T. J. et al. Meta-analysis of the heritability of human traits based on fifty years of twin studies. Nat. Genet. 47, 702\u2013709 (2015).","journal-title":"Nat. Genet."},{"key":"2090_CR20","doi-asserted-by":"publisher","first-page":"m3752","DOI":"10.1136\/bmj.m3752","volume":"371","author":"W-J Chen","year":"2020","unstructured":"Chen, W.-J. et al. Rethinking monogenic neurological diseases. BMJ 371, m3752 (2020).","journal-title":"BMJ"},{"key":"2090_CR21","doi-asserted-by":"publisher","first-page":"eaap8757","DOI":"10.1126\/science.aap8757","volume":"360","author":"The Brainstorm Consortium.","year":"2018","unstructured":"The Brainstorm Consortium. Analysis of shared heritability in common disorders of the brain. Science 360, eaap8757 (2018).","journal-title":"Science"},{"key":"2090_CR22","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1002\/wps.21034","volume":"22","author":"OA Andreassen","year":"2023","unstructured":"Andreassen, O. A., Hindley, G. F. L., Frei, O. & Smeland, O. B. New insights from the last decade of research in psychiatric genetics: discoveries, challenges and clinical implications. World Psychiatry 22, 4\u201324 (2023).","journal-title":"World Psychiatry"},{"key":"2090_CR23","doi-asserted-by":"publisher","first-page":"1339","DOI":"10.1038\/s41588-019-0481-0","volume":"51","author":"K Watanabe","year":"2019","unstructured":"Watanabe, K. et al. A global overview of pleiotropy and genetic architecture in complex traits. Nat. Genet. 51, 1339\u20131348 (2019).","journal-title":"Nat. Genet."},{"key":"2090_CR24","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/j.ajhg.2022.12.011","volume":"110","author":"A Abdellaoui","year":"2023","unstructured":"Abdellaoui, A., Yengo, L., Verweij, K. J. H. & Visscher, P. M. 15 years of GWAS discovery: realizing the promise. Am. J. Hum. Genet 110, 179\u2013194 (2023).","journal-title":"Am. J. Hum. Genet"},{"key":"2090_CR25","doi-asserted-by":"publisher","first-page":"1236","DOI":"10.1038\/ng.3406","volume":"47","author":"B Bulik-Sullivan","year":"2015","unstructured":"Bulik-Sullivan, B. et al. An atlas of genetic correlations across human diseases and traits. Nat. Genet. 47, 1236\u20131241 (2015).","journal-title":"Nat. Genet."},{"key":"2090_CR26","doi-asserted-by":"publisher","first-page":"1469","DOI":"10.1016\/j.cell.2019.11.020","volume":"179","author":"Cross-Disorder Group of the Psychiatric Genomics Consortium.","year":"2019","unstructured":"Cross-Disorder Group of the Psychiatric Genomics Consortium. Genomic relationships, novel loci, and pleiotropic mechanisms across eight psychiatric disorders. Cell 179, 1469\u20131482 (2019).","journal-title":"Cell"},{"key":"2090_CR27","doi-asserted-by":"publisher","first-page":"1795","DOI":"10.1038\/s41588-022-01245-2","volume":"54","author":"C Romero","year":"2022","unstructured":"Romero, C. et al. Exploring the genetic overlap between twelve psychiatric disorders. Nat. Genet. 54, 1795\u20131802 (2022).","journal-title":"Nat. Genet."},{"key":"2090_CR28","doi-asserted-by":"publisher","first-page":"548","DOI":"10.1038\/s41588-022-01057-4","volume":"54","author":"AD Grotzinger","year":"2022","unstructured":"Grotzinger, A. D. et al. Genetic architecture of 11 major psychiatric disorders at biobehavioral, functional genomic and molecular genetic levels of analysis. Nat. Genet. 54, 548\u2013559 (2022).","journal-title":"Nat. Genet."},{"key":"2090_CR29","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/j.neurobiolaging.2023.03.004","volume":"127","author":"DP Wightman","year":"2023","unstructured":"Wightman, D. P. et al. The genetic overlap between Alzheimer\u2019s disease, amyotrophic lateral sclerosis, Lewy body dementia, and Parkinson\u2019s disease. Neurobiol. Aging 127, 99\u2013112 (2023).","journal-title":"Neurobiol. Aging"},{"key":"2090_CR30","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-31873-5","volume":"13","author":"TS Wingo","year":"2022","unstructured":"Wingo, T. S. et al. Shared mechanisms across the major psychiatric and neurodegenerative diseases. Nat. Commun. 13, 4314 (2022).","journal-title":"Nat. Commun."},{"key":"2090_CR31","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-10310-0","volume":"10","author":"O Frei","year":"2019","unstructured":"Frei, O. et al. Bivariate causal mixture model quantifies polygenic overlap between complex traits beyond genetic correlation. Nat. Commun. 10, 2417 (2019).","journal-title":"Nat. Commun."},{"key":"2090_CR32","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1038\/s41588-022-01017-y","volume":"54","author":"J Werme","year":"2022","unstructured":"Werme, J., van der Sluis, S., Posthuma, D. & de Leeuw, C. A. An integrated framework for local genetic correlation analysis. Nat. Genet. 54, 274\u2013282 (2022).","journal-title":"Nat. Genet."},{"key":"2090_CR33","doi-asserted-by":"publisher","first-page":"833","DOI":"10.1176\/appi.ajp.21101051","volume":"179","author":"G Hindley","year":"2022","unstructured":"Hindley, G. et al. Charting the landscape of genetic overlap between mental disorders and related traits beyond genetic correlation. Am. J. Psychiatry 179, 833\u2013843 (2022).","journal-title":"Am. J. Psychiatry"},{"key":"2090_CR34","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1093\/brain\/awab267","volume":"145","author":"S Bahrami","year":"2022","unstructured":"Bahrami, S. et al. Dissecting the shared genetic basis of migraine and mental disorders using novel statistical tools. Brain 145, 142\u2013153 (2022).","journal-title":"Brain"},{"key":"2090_CR35","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1016\/j.biopsych.2020.01.026","volume":"89","author":"OB Smeland","year":"2021","unstructured":"Smeland, O. B. et al. Genome-wide association analysis of Parkinson\u2019s disease and schizophrenia reveals shared genetic architecture and identifies novel risk loci. Biol. Psychiatry 89, 227\u2013235 (2021).","journal-title":"Biol. Psychiatry"},{"key":"2090_CR36","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1038\/ng.3570","volume":"48","author":"JK Pickrell","year":"2016","unstructured":"Pickrell, J. K. et al. Detection and interpretation of shared genetic influences on 42 human traits. Nat. Genet. 48, 709\u2013717 (2016).","journal-title":"Nat. Genet."},{"key":"2090_CR37","doi-asserted-by":"publisher","first-page":"3392","DOI":"10.1093\/brain\/awad038","volume":"146","author":"N Karadag","year":"2023","unstructured":"Karadag, N. et al. Identification of novel genomic risk loci shared between common epilepsies and psychiatric disorders. Brain 146, 3392\u20133403 (2023).","journal-title":"Brain"},{"key":"2090_CR38","doi-asserted-by":"publisher","first-page":"e3002058","DOI":"10.1371\/journal.pbio.3002058","volume":"21","author":"Y Zeighami","year":"2023","unstructured":"Zeighami, Y. et al. A comparison of anatomic and cellular transcriptome structures across 40 human brain diseases. PLoS Biol. 21, e3002058 (2023).","journal-title":"PLoS Biol."},{"key":"2090_CR39","doi-asserted-by":"publisher","first-page":"e1008612","DOI":"10.1371\/journal.pgen.1008612","volume":"16","author":"D Holland","year":"2020","unstructured":"Holland, D. et al. Beyond SNP heritability: polygenicity and discoverability of phenotypes estimated with a univariate Gaussian mixture model. PLoS Genet. 16, e1008612 (2020).","journal-title":"PLoS Genet."},{"key":"2090_CR40","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1038\/s41588-018-0279-5","volume":"51","author":"D Speed","year":"2019","unstructured":"Speed, D. & Balding, D. J. SumHer better estimates the SNP heritability of complex traits from summary statistics. Nat. Genet. 51, 277\u2013284 (2019).","journal-title":"Nat. Genet."},{"key":"2090_CR41","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1038\/ng.3211","volume":"47","author":"BK Bulik-Sullivan","year":"2015","unstructured":"Bulik-Sullivan, B. K. et al. LD Score regression distinguishes confounding from polygenicity in genome-wide association studies. Nat. Genet. 47, 291\u2013295 (2015).","journal-title":"Nat. Genet."},{"key":"2090_CR42","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-21446-3","volume":"12","author":"J Zeng","year":"2021","unstructured":"Zeng, J. et al. Widespread signatures of natural selection across human complex traits and functional genomic categories. Nat. Commun. 12, 1164 (2021).","journal-title":"Nat. Commun."},{"key":"2090_CR43","doi-asserted-by":"publisher","first-page":"704","DOI":"10.1038\/s41586-022-05275-y","volume":"610","author":"L Yengo","year":"2022","unstructured":"Yengo, L. et al. A saturated map of common genetic variants associated with human height. Nature 610, 704\u2013712 (2022).","journal-title":"Nature"},{"key":"2090_CR44","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-017-01261-5","volume":"8","author":"K Watanabe","year":"2017","unstructured":"Watanabe, K., Taskesen, E., van Bochoven, A. & Posthuma, D. Functional mapping and annotation of genetic associations with FUMA. Nat. Commun. 8, 1826 (2017).","journal-title":"Nat. Commun."},{"key":"2090_CR45","doi-asserted-by":"publisher","first-page":"e1004219","DOI":"10.1371\/journal.pcbi.1004219","volume":"11","author":"CA de Leeuw","year":"2015","unstructured":"de Leeuw, C. A., Mooij, J. M., Heskes, T. & Posthuma, D. MAGMA: generalized gene-set analysis of GWAS data. PLoS Comput. Biol. 11, e1004219 (2015).","journal-title":"PLoS Comput. Biol."},{"key":"2090_CR46","first-page":"A42","volume":"3","author":"CW Colton","year":"2006","unstructured":"Colton, C. W. & Manderscheid, R. W. Congruencies in increased mortality rates, years of potential life lost, and causes of death among public mental health clients in eight states. Prev. Chronic Dis. 3, A42 (2006).","journal-title":"Prev. Chronic Dis."},{"key":"2090_CR47","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1038\/s41562-019-0566-x","volume":"3","author":"AD Grotzinger","year":"2019","unstructured":"Grotzinger, A. D. et al. Genomic structural equation modelling provides insights into the multivariate genetic architecture of complex traits. Nat. Hum. Behav. 3, 513\u2013525 (2019).","journal-title":"Nat. Hum. Behav."},{"key":"2090_CR48","doi-asserted-by":"publisher","first-page":"413","DOI":"10.1146\/annurev.neuro.31.060407.125606","volume":"32","author":"PL Strick","year":"2009","unstructured":"Strick, P. L., Dum, R. P. & Fiez, J. A. Cerebellum and nonmotor function. Annu. Rev. Neurosci. 32, 413\u2013434 (2009).","journal-title":"Annu. Rev. Neurosci."},{"key":"2090_CR49","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1038\/s41583-019-0152-2","volume":"20","author":"A Sathyanesan","year":"2019","unstructured":"Sathyanesan, A. et al. Emerging connections between cerebellar development, behaviour and complex brain disorders. Nat. Rev. Neurosci. 20, 298\u2013313 (2019).","journal-title":"Nat. Rev. Neurosci."},{"key":"2090_CR50","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1038\/s41582-020-0364-0","volume":"16","author":"OB Smeland","year":"2020","unstructured":"Smeland, O. B., Frei, O., Dale, A. M. & Andreassen, O. A. The polygenic architecture of schizophrenia\u2014rethinking pathogenesis and nosology. Nat. Rev. Neurol. 16, 366\u2013379 (2020).","journal-title":"Nat. Rev. Neurol."},{"key":"2090_CR51","doi-asserted-by":"publisher","first-page":"2716","DOI":"10.1038\/s41380-023-02076-1","volume":"28","author":"JC Lambert","year":"2023","unstructured":"Lambert, J. C., Ramirez, A., Grenier-Boley, B. & Bellenguez, C. Step by step: towards a better understanding of the genetic architecture of Alzheimer\u2019s disease. Mol. Psychiatry 28, 2716\u20132727 (2023).","journal-title":"Mol. Psychiatry"},{"key":"2090_CR52","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1038\/s43586-021-00056-9","volume":"1","author":"E Uffelmann","year":"2021","unstructured":"Uffelmann, E. et al. Genome-wide association studies. Nat. Rev. Methods Prim. 1, 59 (2021).","journal-title":"Nat. Rev. Methods Prim."},{"key":"2090_CR53","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1038\/s41582-024-00988-2","volume":"20","author":"RJ Jackson","year":"2024","unstructured":"Jackson, R. J., Hyman, B. T. & Serrano-Pozo, A. Multifaceted roles of APOE in Alzheimer disease. Nat. Rev. Neurol. 20, 457\u2013474 (2024).","journal-title":"Nat. Rev. Neurol."},{"key":"2090_CR54","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1001\/jamaneurol.2021.4781","volume":"79","author":"C Liao","year":"2022","unstructured":"Liao, C. et al. Association of essential tremor with novel risk loci: a genome-wide association study and meta-analysis. JAMA Neurol. 79, 185\u2013193 (2022).","journal-title":"JAMA Neurol."},{"key":"2090_CR55","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1038\/s41588-021-00990-0","volume":"54","author":"H Hautakangas","year":"2022","unstructured":"Hautakangas, H. et al. Genome-wide analysis of 102,084 migraine cases identifies 123 risk loci and subtype-specific risk alleles. Nat. Genet. 54, 152\u2013160 (2022).","journal-title":"Nat. Genet."},{"key":"2090_CR56","doi-asserted-by":"publisher","first-page":"954","DOI":"10.1038\/s41593-021-00860-2","volume":"24","author":"DF Levey","year":"2021","unstructured":"Levey, D. F. et al. Bi-ancestral depression GWAS in the Million Veteran Program and meta-analysis in >1.2 million individuals highlight new therapeutic directions. Nat. Neurosci. 24, 954\u2013963 (2021).","journal-title":"Nat. Neurosci."},{"key":"2090_CR57","doi-asserted-by":"publisher","first-page":"1091","DOI":"10.1016\/S1474-4422(19)30320-5","volume":"18","author":"MA Nalls","year":"2019","unstructured":"Nalls, M. A. et al. Identification of novel risk loci, causal insights, and heritable risk for Parkinson\u2019s disease: a meta-analysis of genome-wide association studies. Lancet Neurol. 18, 1091\u20131102 (2019).","journal-title":"Lancet Neurol."},{"key":"2090_CR58","doi-asserted-by":"publisher","first-page":"1505","DOI":"10.1016\/S0140-6736(20)32342-4","volume":"397","author":"M Ashina","year":"2021","unstructured":"Ashina, M. et al. Migraine: integrated approaches to clinical management and emerging treatments. Lancet 397, 1505\u20131518 (2021).","journal-title":"Lancet"},{"key":"2090_CR59","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-17368-1","volume":"11","author":"D van der Meer","year":"2020","unstructured":"van der Meer, D. et al. Understanding the genetic determinants of the brain with MOSTest. Nat. Commun. 11, 3512 (2020).","journal-title":"Nat. Commun."},{"key":"2090_CR60","doi-asserted-by":"publisher","first-page":"1584","DOI":"10.1038\/s41562-023-01630-9","volume":"7","author":"G Hindley","year":"2023","unstructured":"Hindley, G. et al. Multivariate genetic analysis of personality and cognitive traits reveals abundant pleiotropy. Nat. Hum. Behav. 7, 1584\u20131600 (2023).","journal-title":"Nat. Hum. Behav."},{"key":"2090_CR61","doi-asserted-by":"publisher","first-page":"1207","DOI":"10.1038\/s41588-019-0439-2","volume":"51","author":"HJ Watson","year":"2019","unstructured":"Watson, H. J. et al. Genome-wide association study identifies eight risk loci and implicates metabo-psychiatric origins for anorexia nervosa. Nat. Genet. 51, 1207\u20131214 (2019).","journal-title":"Nat. Genet."},{"key":"2090_CR62","doi-asserted-by":"publisher","first-page":"3292","DOI":"10.1038\/s41380-019-0559-1","volume":"25","author":"KL Purves","year":"2020","unstructured":"Purves, K. L. et al. A major role for common genetic variation in anxiety disorders. Mol. Psychiatry 25, 3292\u20133303 (2020).","journal-title":"Mol. Psychiatry"},{"key":"2090_CR63","doi-asserted-by":"crossref","unstructured":"Demontis, D. et al. Genome-wide analyses of ADHD identify 27 risk loci, refine the genetic architecture and implicate several cognitive domains. Nat. Genet. 55, 198\u2013208 (2023).","DOI":"10.1016\/j.euroneuro.2022.07.018"},{"key":"2090_CR64","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1038\/s41588-019-0344-8","volume":"51","author":"J Grove","year":"2019","unstructured":"Grove, J. et al. Identification of common genetic risk variants for autism spectrum disorder. Nat. Genet. 51, 431\u2013444 (2019).","journal-title":"Nat. Genet."},{"key":"2090_CR65","doi-asserted-by":"publisher","first-page":"817","DOI":"10.1038\/s41588-021-00857-4","volume":"53","author":"N Mullins","year":"2021","unstructured":"Mullins, N. et al. Genome-wide association study of more than 40,000 bipolar disorder cases provides new insights into the underlying biology. Nat. Genet. 53, 817\u2013829 (2021).","journal-title":"Nat. Genet."},{"key":"2090_CR66","doi-asserted-by":"crossref","unstructured":"International Obsessive Compulsive Disorder Foundation Genetics Collaborative (IOCDF-GC) and OCD Collaborative Genetics Association Studies (OCGAS). Revealing the complex genetic architecture of obsessive-compulsive disorder using meta-analysis. Mol. Psychiatry 23, 1181\u20131188 (2018).","DOI":"10.1038\/mp.2017.154"},{"key":"2090_CR67","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-12576-w","volume":"10","author":"CM Nievergelt","year":"2019","unstructured":"Nievergelt, C. M. et al. International meta-analysis of PTSD genome-wide association studies identifies sex- and ancestry-specific genetic risk loci. Nat. Commun. 10, 4558 (2019).","journal-title":"Nat. Commun."},{"key":"2090_CR68","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1038\/s41586-022-04434-5","volume":"604","author":"V Trubetskoy","year":"2022","unstructured":"Trubetskoy, V. et al. Mapping genomic loci implicates genes and synaptic biology in schizophrenia. Nature 604, 502\u2013508 (2022).","journal-title":"Nature"},{"key":"2090_CR69","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1176\/appi.ajp.2018.18070857","volume":"176","author":"D Yu","year":"2019","unstructured":"Yu, D. et al. Interrogating the genetic determinants of Tourette's syndrome and other tic disorders through genome-wide association studies. Am. J. Psychiatry 176, 217\u2013227 (2019).","journal-title":"Am. J. Psychiatry"},{"key":"2090_CR70","doi-asserted-by":"publisher","first-page":"1276","DOI":"10.1038\/s41588-021-00921-z","volume":"53","author":"DP Wightman","year":"2021","unstructured":"Wightman, D. P. et al. A genome-wide association study with 1,126,563 individuals identifies new risk loci for Alzheimer's disease. Nat. Genet. 53, 1276\u20131282 (2021).","journal-title":"Nat. Genet"},{"key":"2090_CR71","doi-asserted-by":"publisher","first-page":"1636","DOI":"10.1038\/s41588-021-00973-1","volume":"53","author":"W van Rheenen","year":"2021","unstructured":"van Rheenen, W. et al. Common and rare variant association analyses in amyotrophic lateral sclerosis identify 15 risk loci with distinct genetic architectures and neuron-specific biology. Nat. Genet. 53, 1636\u20131648 (2021).","journal-title":"Nat. Genet."},{"key":"2090_CR72","doi-asserted-by":"publisher","first-page":"1471","DOI":"10.1038\/s41588-023-01485-w","volume":"55","author":"International League Against Epilepsy Consortium on Complex Epilepsies.","year":"2023","unstructured":"International League Against Epilepsy Consortium on Complex Epilepsies. GWAS meta-analysis of over 29,000 people with epilepsy identifies 26 risk loci and subtype-specific genetic architecture. Nat. Genet. 55, 1471\u20131482 (2023).","journal-title":"Nat. Genet."},{"key":"2090_CR73","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1038\/s41588-021-00785-3","volume":"53","author":"R Chia","year":"2021","unstructured":"Chia, R. et al. Genome sequencing analysis identifies new loci associated with Lewy body dementia and provides insights into its genetic architecture. Nat. Genet. 53, 294\u2013303 (2021).","journal-title":"Nat. Genet."},{"key":"2090_CR74","unstructured":"International Multiple Sclerosis Genetics Consortium. Multiple sclerosis genomic map implicates peripheral immune cells and microglia in susceptibility. Science 365, eaav7188 (2019)."},{"key":"2090_CR75","doi-asserted-by":"publisher","first-page":"989","DOI":"10.1038\/ng.3043","volume":"46","author":"MA Nalls","year":"2014","unstructured":"Nalls, M. A. et al. Large-scale meta-analysis of genome-wide association data identifies six new risk loci for Parkinson's disease. Nat. Genet. 46, 989\u2013993 (2014).","journal-title":"Nat. Genet."},{"key":"2090_CR76","doi-asserted-by":"publisher","first-page":"1511","DOI":"10.1038\/ng.3955","volume":"49","author":"D Chang","year":"2017","unstructured":"Chang, D. et al. A meta-analysis of genome-wide association studies identifies 17 new Parkinson's disease risk loci. Nat. Genet. 49, 1511\u20131516 (2017).","journal-title":"Nat. Genet."},{"key":"2090_CR77","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1038\/s41586-022-05165-3","volume":"611","author":"A Mishra","year":"2022","unstructured":"Mishra, A. et al. Stroke genetics informs drug discovery and risk prediction across ancestries. Nature 611, 115\u2013123 (2022).","journal-title":"Nature"},{"key":"2090_CR78","doi-asserted-by":"publisher","first-page":"912","DOI":"10.1038\/s41588-018-0152-6","volume":"50","author":"JE Savage","year":"2018","unstructured":"Savage, J. E. et al. Genome-wide association meta-analysis in 269,867 individuals identifies new genetic and functional links to intelligence. Nat. Genet. 50, 912\u2013919 (2018).","journal-title":"Nat. Genet."},{"key":"2090_CR79","doi-asserted-by":"publisher","first-page":"957","DOI":"10.1038\/s41588-019-0407-x","volume":"51","author":"M Wuttke","year":"2019","unstructured":"Wuttke, M. et al. A catalog of genetic loci associated with kidney function from analyses of a million individuals. Nat. Genet. 51, 957\u2013972 (2019).","journal-title":"Nat. Genet."},{"key":"2090_CR80","doi-asserted-by":"publisher","first-page":"1385","DOI":"10.1038\/ng.3913","volume":"49","author":"CP Nelson","year":"2017","unstructured":"Nelson, C. P. et al. Association analyses based on false discovery rate implicate new loci for coronary artery disease. Nat. Genet. 49, 1385\u20131391 (2017).","journal-title":"Nat. Genet."},{"key":"2090_CR81","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1038\/ng.3760","volume":"49","author":"KM de Lange","year":"2017","unstructured":"de Lange, K. M. et al. Genome-wide association study implicates immune activation of multiple integrin genes in inflammatory bowel disease. Nat. Genet. 49, 256\u2013261 (2017).","journal-title":"Nat. Genet."},{"key":"2090_CR82","doi-asserted-by":"publisher","first-page":"1505","DOI":"10.1038\/s41588-018-0241-6","volume":"50","author":"A Mahajan","year":"2018","unstructured":"Mahajan, A. et al. Fine-mapping type 2 diabetes loci to single-variant resolution using high-density imputation and islet-specific epigenome maps. Nat. Genet. 50, 1505\u20131513 (2018).","journal-title":"Nat. Genet."},{"key":"2090_CR83","doi-asserted-by":"publisher","unstructured":"Gadin, J.R., Zetterberg, R., Meijsen, J. & Schork, A.J. Cleansumstats: converting GWAS sumstats to a common format to facilitate downstream applications. Zenodo https:\/\/doi.org\/10.5281\/zenodo.7441266 (2023).","DOI":"10.5281\/zenodo.7441266"},{"key":"2090_CR84","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1038\/nature15393","volume":"526","author":"The 1000 Genomes Project Consortium.","year":"2015","unstructured":"The 1000 Genomes Project Consortium. A global reference for human genetic variation. Nature 526, 68\u201374 (2015).","journal-title":"Nature"},{"key":"2090_CR85","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1146\/annurev.genom.7.080505.115623","volume":"7","author":"GA Maston","year":"2006","unstructured":"Maston, G. A., Evans, S. K. & Green, M. R. Transcriptional regulatory elements in the human genome. Annu. Rev. Genomics Hum. Genet. 7, 29\u201359 (2006).","journal-title":"Annu. Rev. Genomics Hum. Genet."},{"key":"2090_CR86","doi-asserted-by":"publisher","first-page":"1.30.1","DOI":"10.1002\/cpbi.5","volume":"54","author":"G Stelzer","year":"2016","unstructured":"Stelzer, G. et al. The GeneCards Suite: from gene data mining to disease genome sequence analyses. Curr. Protoc. Bioinformatics 54, 1.30.1\u20131.30.33 (2016).","journal-title":"Curr. Protoc. Bioinformatics"},{"key":"2090_CR87","doi-asserted-by":"publisher","first-page":"1739","DOI":"10.1093\/bioinformatics\/btr260","volume":"27","author":"A Liberzon","year":"2011","unstructured":"Liberzon, A. et al. Molecular signatures database (MSigDB) 3.0. Bioinformatics 27, 1739\u20131740 (2011).","journal-title":"Bioinformatics"},{"key":"2090_CR88","doi-asserted-by":"publisher","first-page":"580","DOI":"10.1038\/ng.2653","volume":"45","author":"The GTEx Consortium.","year":"2013","unstructured":"The GTEx Consortium. The Genotype\u2013Tissue Expression (GTEx) project. Nat. Genet. 45, 580\u2013585 (2013).","journal-title":"Nat. Genet."},{"key":"2090_CR89","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-11181-1","volume":"10","author":"K Watanabe","year":"2019","unstructured":"Watanabe, K., Umicevic Mirkov, M., de Leeuw, C. A., van den Heuvel, M. P. & Posthuma, D. Genetic mapping of cell type specificity for complex traits. Nat. Commun. 10, 3222 (2019).","journal-title":"Nat. Commun."},{"key":"2090_CR90","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1038\/s41586-019-1506-7","volume":"573","author":"RD Hodge","year":"2019","unstructured":"Hodge, R. D. et al. Conserved cell types with divergent features in human versus mouse cortex. Nature 573, 61\u201368 (2019).","journal-title":"Nature"},{"key":"2090_CR91","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1038\/nmeth.4407","volume":"14","author":"N Habib","year":"2017","unstructured":"Habib, N. et al. Massively parallel single-nucleus RNA-seq with DroNc-seq. Nat. Methods 14, 955\u2013958 (2017).","journal-title":"Nat. Methods"},{"key":"2090_CR92","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1038\/nature25980","volume":"555","author":"S Zhong","year":"2018","unstructured":"Zhong, S. et al. A single-cell RNA-seq survey of the developmental landscape of the human prefrontal cortex. Nature 555, 524\u2013528 (2018).","journal-title":"Nature"},{"key":"2090_CR93","doi-asserted-by":"publisher","first-page":"7285","DOI":"10.1073\/pnas.1507125112","volume":"112","author":"S Darmanis","year":"2015","unstructured":"Darmanis, S. et al. A survey of human brain transcriptome diversity at the single cell level. Proc. Natl Acad. Sci. USA 112, 7285\u20137290 (2015).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2090_CR94","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-16546-4","volume":"7","author":"H Hochgerner","year":"2017","unstructured":"Hochgerner, H. et al. STRT-seq-2i: dual-index 5\u2019 single cell and nucleus RNA-seq on an addressable microwell array. Sci. Rep. 7, 16327 (2017).","journal-title":"Sci. Rep."},{"key":"2090_CR95","doi-asserted-by":"publisher","first-page":"566","DOI":"10.1016\/j.cell.2016.09.027","volume":"167","author":"G La Manno","year":"2016","unstructured":"La Manno, G. et al. Molecular diversity of midbrain development in mouse, human, and stem cells. Cell 167, 566\u2013580 (2016).","journal-title":"Cell"},{"key":"2090_CR96","doi-asserted-by":"publisher","first-page":"eaat8464","DOI":"10.1126\/science.aat8464","volume":"362","author":"D Wang","year":"2018","unstructured":"Wang, D. et al. Comprehensive functional genomic resource and integrative model for the human brain. Science 362, eaat8464 (2018).","journal-title":"Science"},{"key":"2090_CR97","doi-asserted-by":"publisher","first-page":"4428","DOI":"10.1016\/j.cell.2022.09.039","volume":"185","author":"CA Herring","year":"2022","unstructured":"Herring, C. A. et al. Human prefrontal cortex gene regulatory dynamics from gestation to adulthood at single-cell resolution. Cell 185, 4428\u20134447 (2022).","journal-title":"Cell"},{"key":"2090_CR98","doi-asserted-by":"publisher","first-page":"1165","DOI":"10.1214\/aos\/1013699998","volume":"29","author":"Y Benjamini","year":"2001","unstructured":"Benjamini, Y. & Yekutieli, D. The control of the false discovery rate in multiple testing under dependency. Ann. Stat. 29, 1165\u20131188 (2001).","journal-title":"Ann. Stat."}],"container-title":["Nature Neuroscience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41593-025-02090-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41593-025-02090-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41593-025-02090-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,2]],"date-time":"2025-12-02T23:02:01Z","timestamp":1764716521000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41593-025-02090-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,11]]},"references-count":98,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2025,12]]}},"alternative-id":["2090"],"URL":"https:\/\/doi.org\/10.1038\/s41593-025-02090-2","relation":{},"ISSN":["1097-6256","1546-1726"],"issn-type":[{"value":"1097-6256","type":"print"},{"value":"1546-1726","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,11]]},"assertion":[{"value":"30 May 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 September 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 November 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"O.A.A. has received speaker\u2019s honoraria from Lundbeck, Sunovion, Takeda and Janssen and is a consultant for Cortechs.ai. A.M.D. is a founder of and holds equity interest in CorTechs Labs and serves on its scientific advisory board. He is also a member of the Scientific Advisory Board of Healthlytix and receives research funding from General Electric Healthcare. The terms of these arrangements have been reviewed and approved by the University of California, San Diego, in accordance with its conflict-of-interest policies. T.E. is a consultant to Cumulus and Sumitomo Pharma and received speaker\u2019s honoraria from Lundbeck and Janssen Cilag. The other authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}