# Dissecting the Shared Genetic Architecture of Suicide Attempt, Psychiatric Disorders, and Known Risk Factors

Niamh Mullins, Jooeun Kang, Adrian I. Campos, Jonathan R.I. Coleman, Alexis C. Edwards, Hanga Galfalvy, Daniel F. Levey, Adriana Lori, Andrey Shabalin, Anna Starnawska, et al.

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## To cite this version:

Niamh Mullins, Jooeun Kang, Adrian I. Campos, Jonathan R.I. Coleman, Alexis C. Edwards, et al. Dissecting the Shared Genetic Architecture of Suicide Attempt, Psychiatric Disorders, and Known Risk Factors. Biological Psychiatry, 2022, 91 (3), pp.313 - 327. 10.1016/j.biopsych.2021.05.029. inserm-05086841

HAL Id: inserm-05086841

https://inserm.hal.science/inserm-05086841v1

Submitted on 27 May 2025

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Dissecting the Shared Genetic Architecture of
Suicide Attempt, Psychiatric Disorders, and
Known Risk Factors

Niamh Mullins, JooEun Kang, Adrian I. Campos, Jonathan R.I. Coleman, Alexis C. Edwards, Hanga Galfalvy, Daniel F. Levey, Adriana Lori, Andrey Shabalin, Anna Starnawska, Mei-Hsin Su, Hunna J. Watson, Mark Adams, Swapnil Awasthi, Michael Gandal, Jonathan D. Hafferty, Akitoyo Hishimoto, Minsoo Kim, Satoshi Okazaki, Ikuo Otsuka, Stephan Ripke, Erin B. Ware, Andrew W. Bergen, Wade H. Berrettini, Martin Bohus, Harry Brandt, Xiao Chang, Wei J. Chen, Hsi-Chung Chen, Steven Crawford, Scott Crow, Emily DiBlasi, Philibert Duriez, Fernando Fernández-Aranda, Manfred M. Fichter, Steven Gallinger, Stephen J. Glatt, Philip Gorwood, Yiran Guo, Hakon Hakonarson, Katherine A. Halmi, Hai-Gwo Hwu, Sonia Jain, Stéphane Jamain, Susana Jiménez-Murcia, Craig Johnson, Allan S. Kaplan, Walter H. Kaye, Pamela K. Keel, James L. Kennedy, Kelly L. Klump, Dong Li, Shih-Cheng Liao, Klaus Lieb, Lisa Lilenfeld, Chih-Min Liu, Pierre J. Magistretti, Christian R. Marshall, James E. Mitchell, Eric T. Monson, Richard M. Myers, Dalila Pinto, Abigail Powers, Nicolas Ramoz, Stefan Roepke, Vsevolod Rozanov, Stephen W. Scherer, Christian Schmahl, Marcus Sokolowski, Michael Strober, Laura M. Thornton, Janet Treasure, Ming T. Tsuang, Stephanie H. Witt, D. Blake Woodside, Zeynep Yilmaz, Lea Zillich, Rolf Adolfsson, Ingrid Agartz, Tracy M. Air, Martin Alda, Lars Alfredsson, Ole A. Andreassen, Adebayo Anjorin, Vivek Appadurai, María Soler Artigas, Sandra Van der Auwera, M. Helena Azevedo, Nicholas Bass, Claiton H.D. Bau, Bernhard T. Baune, Frank Bellivier, Klaus Berger, Joanna M. Biernacka, Tim B. Bigdeli, Elisabeth B. Binder, Michael Boehnke, Marco P. Boks, Rosa Bosch, David L. Braff, Richard Bryant, Monika Budde, Enda M. Byrne, Wiepke Cahn, Miguel Casas, Enrique Castelao, Jorge A. Cervilla, Boris Chaumette, Sven Cichon, Aiden Corvin, Nicholas Craddock, David Craig, Franziska Degenhardt, Srdjan Djurovic, Howard J. Edenberg, Ayman H. Fanous, Jerome C. Foo, Andreas J. Forstner, Mark Frye, Janice M. Fullerton, Justine M. Gatt, Pablo V. Gejman, Ina Giegling, Hans J. Grabe, Melissa J. Green, Eugenio H. Grevet, Maria Grigoreiu-Serbanescu, Blanca Gutierrez, Jose Guzman-Parra, Steven P. Hamilton, Marian L. Hamshere, Annette Hartmann, Joanna Hauser, Stefanie Heilmann-Heimbach, Per Hoffmann, Marcus Ising, Ian Jones, Lisa A. Jones, Lina Jonsson, René S. Kahn, John R. Kelsoe, Kenneth S. Kendler, Stefan Kloiber, Karestan C. Koenen, Manolis Kogevinas, Bettina Konte, Marie-Odile Krebs, Mikael Landén, Jacob Lawrence, Marion Leboyer, Phil H. Lee, Douglas F. Levinson, Calwing Liao, Jolanta Lissowska, Susanne Lucae, Fermin Mayoral, Susan L. McElroy, Patrick McGrath, Peter McGuffin, Andrew McQuillin, Sarah E. Medland, Divya Mehta, Ingrid Melle, Yuri Milaneschi, Philip B. Mitchell, Esther Molina, Gunnar Morken, Preben Bo Mortensen, Bertram Müller-Myhsok, Caroline Nievergelt, Vishwajit Nimgaonkar, Markus M. Nöthen, Michael C. O'Donovan, Roel A. Ophoff, Michael J. Owen, Carlos Pato, Michele T. Pato, Brenda W.J.H. Penninx, Jonathan Pimm, Giorgio Pistis, James B. Potash, Robert A. Power, Martin Preisig, Digby Quested, Josep Antoni Ramos-Quiroga, Andreas Reif, Marta Ribasés, Vanesa Richarte, Marcella Rietschel, Margarita Rivera, Andrea Roberts, Gloria Roberts, Guy A. Rouleau, Diego L. Rovaris, Dan Rujescu, Cristina Sánchez-Mora, Alan R. Sanders, Peter R. Schofield, Thomas G. Schulze, Laura J. Scott, Alessandro Serretti, Jianxin Shi, Stanley I. Shyn, Lea Sirignano, Pamela Sklar, Olav B. Smeland, Jordan W. Smoller, Edmund J.S. Sonuga-Barke, Gianfranco Spalletta, John S. Strauss, Beata Świątkowska, Maciej Trzaskowski, Gustavo Turecki, Laura Vilar-Ribó, John B. Vincent, Henry Völzke,
James T.R. Walters, Cynthia Shannon Weickert, Thomas W. Weickert, Myrna M. Weissman, Leanne M. Williams, Naomi R. Wray, Clement C. Zai, Allison E. Ashley-Koch, Jean C. Beckham, Elizabeth R. Hauser, Michael A. Hauser, Nathan A. Kimbrel, Jennifer H. Lindquist, Benjamin McMahon, David W. Oslin, Xuejun Qin, Major Depressive Disorder Working Group of the Psychiatric Genomics Consortium, Bipolar Disorder Working Group of the Psychiatric Genomics Consortium, Eating Disorders Working Group of the Psychiatric Genomics Consortium, German Borderline Genomics Consortium, MVP Suicide Exemplar Workgroup, VA Million Veteran Program, Esben Agerbo, Anders D. Børglum, Gerome Breen, Annette Erlangsen, Tõnu Esko, Joel Gelernter, David M. Hougaard, Ronald C. Kessler, Henry R. Kranzler, Qingqin S. Li, Nicholas G. Martin, Andrew M. McIntosh, Ole Mors, Merete Nordentoft, Catherine M. Olsen, David Porteous, Robert J. Ursano, Danuta Wasserman, Thomas Werge, David C. Whiteman, Cynthia M. Bulik, Hilary Coon, Ditte Demontis, Anna R. Docherty, Po-Hsiu Kuo, Kathryn M. Lewis, J. John Mann, Miguel E. Rentería, Daniel J. Smith, Eli A. Stahl, Murray B. Stein, Fabian Streit, Virginia Willour, and Douglas M. Ruderfer

## ABSTRACT

**BACKGROUND:** Suicide is a leading cause of death worldwide, and nonfatal suicide attempts, which occur far more frequently, are a major source of disability and social and economic burden. Both have substantial genetic etiology, which is partially shared and partially distinct from that of related psychiatric disorders.

**METHODS:** We conducted a genome-wide association study (GWAS) of 29,782 suicide attempt (SA) cases and 519,961 controls in the International Suicide Genetics Consortium (ISGC). The GWAS of SA was conditioned on psychiatric disorders using GWAS summary statistics via multitrait-based conditional and joint analysis, to remove genetic effects on SA mediated by psychiatric disorders. We investigated the shared and divergent genetic architectures of SA, psychiatric disorders, and other known risk factors.

**RESULTS:** Two loci reached genome-wide significance for SA: the major histocompatibility complex and an intergenic locus on chromosome 7, the latter of which remained associated with SA after conditioning on psychiatric disorders and replicated in an independent cohort from the Million Veteran Program. This locus has been implicated in risk-taking behavior, smoking, and insomnia. SA showed strong genetic correlation with psychiatric disorders, particularly major depression, and also with smoking, pain, risk-taking behavior, sleep disturbances, lower educational attainment, reproductive traits, lower socioeconomic status, and poorer general health. After conditioning on psychiatric disorders, the genetic correlations between SA and psychiatric disorders decreased, whereas those with nonpsychiatric traits remained largely unchanged.

**CONCLUSIONS:** Our results identify a risk locus that contributes more strongly to SA than other phenotypes and suggest a shared underlying biology between SA and known risk factors that is not mediated by psychiatric disorders.

https://doi.org/10.1016/j.biopsych.2021.05.029

Suicide is a worldwide public health problem, accounting for almost 800,000 deaths per year (1). Nonfatal suicide attempt (SA), defined as self-injurious behavior with the intent to die, has been estimated to occur over 20 times more frequently and is a major source of disability, reduced quality of life, and social and economic burden (1,2). The lifetime prevalence of SA in adults ranges from 0.5% to 5% worldwide (3). There are several well-established comorbidities and risk factors for SA, with psychiatric illness having the strongest effect on lifetime suicide rates (4,5). However, the vast majority of patients with psychiatric disorders never attempt suicide (6–8). Other major risk factors for SA include prior self-injurious thoughts and behaviors (9), physical illness or disability (10,11), sleep disorders (12–15), family history of psychiatric disorders (16),

substance abuse (17), smoking (18–20), impulsivity (21) and social factors including childhood maltreatment (21), isolation (22), and stressful life events (23).

Both suicide and SA are heritable, with estimates from genetic epidemiology studies ranging from 17% to 55% (24–26). Several genome-wide association studies (GWASs) of SA have reported significant single nucleotide polymorphism (SNP)-heritability estimates of ~4%, indicating an underlying polygenic architecture (27–31). Using polygenic risk scoring or genetic correlation analyses, these studies have also demonstrated shared genetic etiology between SA and psychiatric disorders, with major depressive disorder (MDD) showing the largest genetic overlap (28,29,31). This genetic overlap, along with the high prevalence of MDD in the population (32), make it
<table><caption>**Table 1. Numbers of Cases and Controls for 18 Cohorts in the International Suicide Genetics Consortium**</caption><thead><tr><th>Cohort (Ancestry)</th><th>SA Cases</th><th>Controls</th></tr></thead><tbody><tr><td>Psychiatric Genomics Consortium MDD (EUR)</td><td>1528</td><td>16,626</td></tr><tr><td>Psychiatric Genomics Consortium BIP (EUR)</td><td>3214</td><td>17,642</td></tr><tr><td>Psychiatric Genomics Consortium SCZ (EUR)</td><td>1640</td><td>7112</td></tr><tr><td>Psychiatric Genomics Consortium ED (EUR)</td><td>170</td><td>5070</td></tr><tr><td>Army STARRS (EUR)</td><td>670</td><td>10,637</td></tr><tr><td>German Borderline Genomics Consortium (EUR)</td><td>481</td><td>1653</td></tr><tr><td>UK Biobank (EUR)</td><td>2433</td><td>334,766</td></tr><tr><td>iPSYCH (EUR)</td><td>7003</td><td>52,227</td></tr><tr><td>Janssen (EUR)</td><td>255</td><td>1684</td></tr><tr><td>Yale-Penn (EUR)</td><td>475</td><td>1817</td></tr><tr><td>GISS Ukraine (EUR)</td><td>660</td><td>660</td></tr><tr><td>Columbia University (EUR)</td><td>577</td><td>1233</td></tr><tr><td>Australian Genetics of Depression Study and QSkin Study (EUR)</td><td>2792</td><td>20,193</td></tr><tr><td>University of Utah (EUR)</td><td>4692</td><td>20,702</td></tr><tr><td>Japan (EAS)</td><td>746</td><td>14,049</td></tr><tr><td>CONVERGE (EAS)</td><td>1148</td><td>6515</td></tr><tr><td>Grady Trauma Project (Admixed AA)</td><td>669</td><td>4473</td></tr><tr><td>Yale-Penn (Admixed AA)</td><td>629</td><td>2902</td></tr><tr><td>Total</td><td>29,782</td><td>519,961</td></tr></tbody></table>

AA, African American; Army STARRS, Army Study to Assess Risk and Resilience in Servicemembers; BIP, bipolar disorder; EAS, East Asian; ED, eating disorder; EUR, European; GISS, Genetic Investigation of Suicide and Suicide Attempt; MDD, major depressive disorder; SA, suicide attempt; SCZ, schizophrenia.

cohort. SNPs with a weighted minor allele frequency of <1%, weighted INFO score <0.6, or SNPs present in <80% of total effective sample size were removed from the meta-analysis results. A genome-wide significant locus was defined as the region around a SNP with $p < 5.0 \times 10^{-8}$ with linkage disequilibrium (LD) $r^2 > 0.1$, within a 3000 kb window, based on the LD structure of the Haplotype Reference Consortium European ancestries reference panel (version 1.0) (37).

## Statistical Conditioning on Psychiatric Disorders

The results of the SA-EUR meta-analysis were conditioned on the genetics of MDD using multitrait-based conditional and joint analysis using GWAS summary data (mtCOJO) (38), implemented in the GCTA software package (39). mtCOJO (38) estimates the effect size of a SNP on an outcome trait conditioned on exposure trait(s). Genome-wide significant SNPs for the exposure are used as instruments to estimate the effect of the exposure on the outcome, and this effect is used to perform genome-wide conditioning, yielding conditioned effect sizes and *p* values for the outcome trait. We conditioned SA (outcome) on MDD (exposure), because MDD is the most prevalent psychiatric disorder among individuals who die by suicide (40) and has the highest genetic correlation with SA (28). The SA-EUR GWAS summary statistics were used as the outcome trait, because mtCOJO requires an ancestry-matched LD reference panel and GWAS summary statistics for the exposure trait. The PGC MDD GWAS results (excluding 23andMe) (41) were used as the exposure, and the results

yielded GWAS summary statistics for SA conditioned on MDD (SA-EUR|MDD). mtCOJO is robust to sample overlap between the GWAS of the exposure and outcome. To select SNPs as instruments, independence was defined as SNPs more than 1 megabase apart or with LD $r^2 < 0.05$ based on the 1000 Genomes Project Phase 3 EUR reference panel (42). To obtain at least 10 independent instruments for MDD, the genome-wide significance threshold was adjusted to $p < 5.0 \times 10^{-7}$, leading to 15 SNPs used. In a further sensitivity analysis, GWAS summary statistics for BIP (43) and SCZ (44) were additionally included as exposure traits.

## LD Score Regression

LDSC (34) was used to estimate the phenotypic variance in SA explained by common SNPs (SNP-heritability, $h_{SNP}^2$) from GWAS summary statistics. $h_{SNP}^2$ was calculated on the liability scale assuming a lifetime prevalence of SA in the general population of 2% (middle of the range reported worldwide) (3). The bivariate genetic correlation attributable to genome-wide SNPs ($r_g$) was estimated between the SA-EUR and SA-EUR|MDD GWAS and a range of psychiatric disorders, self-harm ideation, and propensity toward risk-taking behavior, using the largest available GWAS summary statistics (Bonferroni-corrected significance threshold $p < .0042$, adjusting for 12 traits tested). Differences in $r_g$ with SA-EUR versus SA-EUR|MDD were tested for deviation from 0, using the block jack-knife method, implemented in LDSC software (45). The $r_g$s of SA-EUR and SA-EUR|MDD with 768 other nonoverlapping human diseases and traits were calculated on LD Hub (46) (Bonferroni-corrected significance threshold $p < 6.51 \times 10^{-5}$). Traits were precategorized manually into 15 risk factor groups previously ascribed to SA (4,5,10): autoimmune disease, neurologic disease, heart disease, hypertension, diabetes, kidney disease, cancer, alcohol use, smoking, pain, psychiatric, sleep, life stressors, socioeconomic, and education/cognition. There were 259 traits belonging to these categories, and a second reviewer validated the categories assigned to traits and their relevance to SA risk.

## Polygenic Risk Scoring

Polygenic risk scores (PRSs) for SA were tested for association with SA or death by suicide versus controls in 7 target cohorts: PGC MDD, BIP and SCZ, CONVERGE (EAS ancestries), the University of Utah (suicide death cohort), Yale-Penn (AA ancestries), and Grady Trauma Project (AA ancestries). The primary SA GWAS meta-analysis was repeated excluding each cohort in turn, to create independent discovery datasets. PRSs were generated using PRS-CS (47), which uses a Bayesian regression framework to place continuous shrinkage priors on effect sizes of SNPs in the PRS, adaptive to the strength of their association signal in the discovery GWAS, and the LD structure from an external reference panel (47). The 1000 Genomes EUR, EAS, or African reference panels (42) were used to estimate LD between SNPs, as appropriate for each target cohort. PLINK 1.9 (48) was used to weight SNPs by their effect sizes calculated using PRS-CS and sum all SNPs into PRS for each individual in the target cohorts. PRSs were tested for association with case versus control status in the target cohort using a logistic regression model including covariates as per
the GWAS. The amount of phenotypic variance explained by the PRS ($R^2$) was calculated on the liability scale, assuming a lifetime prevalence of SA in the general population of 2% (3). Analyses in the PGC cohorts were repeated using PRSs generated from the SA-EUR|MDD GWAS results, excluding each PGC cohort in turn. Analyses performed are summarized in Table S2 in Supplement 2 (Bonferroni-corrected significance threshold $p < 3.12 \times 10^{-3}$, adjusting for 16 tests).

## RESULTS

### SA Shows Significant SNP-Heritability and PRS Associations

The primary SA GWAS included 29,782 cases and 519,961 controls from 18 cohorts (Table 1). Cases were predominantly of EUR ancestries (90%), with 6% of EAS ancestries and 4% of admixed AA ancestries. Case definition was lifetime SA, with ~20% of cases having died by suicide. The SNP-heritability ($h_{SNP}^2$) SA was 6.8% (SE = 0.005, $p = 2.00 \times 10^{-42}$) on the liability scale. The LDSC intercept was 1.04 (SE = 0.01, $p = 2.84 \times 10^{-4}$), and the attenuation ratio was 0.14 (SE = 0.04), indicating that the majority of GWAS test statistic inflation was due to polygenicity (Figure S1 in Supplement 1). PRSs for SA were tested in 7 target cohorts (Table S2 in Supplement 2). SA PRSs were significantly associated with SA in the PGC MDD, BIP, and SCZ cohorts, with a phenotypic explained variance ($R^2$) of 0.69% ($p = 7.17 \times 10^{-15}$), 0.68% ($p = 8.11 \times 10^{-28}$), and 0.88% ($p = 1.24 \times 10^{-17}$), respectively (liability scale). PRSs for SA were also associated with death by suicide in the University of Utah cohort, explaining slightly more phenotypic variance ($R^2 = 1.08\%$, $p = 9.79 \times 10^{-81}$). The $r_g$ between the University of Utah suicide death GWAS and a meta-analysis of the nonfatal SA cohorts in our study was 0.77 (SE = 0.08, $p = 3.08 \times 10^{-20}$). Examining the performance of SA PRSs across ancestries showed a significant association with SA in the CONVERGE EAS cohort, although with a lower explained variance ($R^2 = 0.25\%$, $p = 3.06 \times 10^{-3}$). Analyses in 2 admixed AA cohorts showed variable results ($R^2 = 0.21\%$, $p = 5.28 \times 10^{-1}$ and $R^2 = 0.58\%$, $p = 3.44 \times 10^{-3}$, respectively) (Table S2 in Supplement 2).

### GWAS of SA Identifies Locus With Stronger Effect on SA Than Psychiatric Disorders

The primary SA GWAS identified 2 genome-wide significant loci ($p < 5 \times 10^{-9}$) (Table S3 in Supplement 2). The most strongly associated locus was in an intergenic region on chromosome 7 (index SNP rs62474683, odds ratio A allele = 1.06 [1.04–1.08], $p = 1.91 \times 10^{-10}$, frequency in cases = 0.52, frequency in controls = 0.50, $I^2$ heterogeneity index = 0%) (forest plot Figure S2 in Supplement 1). The second genome-wide significant locus was in the major histocompatibility complex (MHC) (index SNP rs71557378, odds ratio T allele = 1.10 [1.06–1.13], $p = 1.97 \times 10^{-8}$, frequency in cases = 0.91, frequency in controls = 0.90, $I^2$ heterogeneity index = 46%) (forest plot Figure S3 in Supplement 1). Both loci were also genome-wide significant in the SA-EUR meta-analysis, with the same effect sizes (Table S4 in Supplement 2). In order to identify SA genetic effects not mediated by MDD, we conditioned the SA-EUR GWAS on the genetic effects of MDD via

mtCOJO. After conditioning, only the chromosome 7 locus remained genome-wide significant (index SNP = rs62474683, odds ratio A allele = 1.06 [1.04–1.08], $p = 1.33 \times 10^{-8}$) (Figure 1A). Figures S4 and S5 in Supplement 1 show regional association plots of the loci before and after conditioning. The association of the chromosome 7 index SNP with SA was further replicated in the independent Million Veteran Program cohort (rs62474683, odds ratio A allele = 1.03 [1.01–1.07], $p = 3.27 \times 10^{-3}$), while the index SNP in the MHC was not associated with SA in this cohort (Table S4 in Supplement 2).

Examination of the chromosome 7 locus in published GWAS results using the Open Targets Genetics web portal (49) indicated smaller and nonsignificant effects on all psychiatric disorders (Figure 1B). Additionally, the SA-index SNP has been implicated at genome-wide significance in lifetime smoking index (50) (accounts for duration and amount of smoking) and propensity toward risk-taking behavior (51), although again with smaller effect sizes than on SA (Figure 1B; Tables S5 and S6 in Supplement 2). Pairwise GWAS analysis (see Supplement 1) of the genomic region containing the chromosome 7 locus suggested the existence of a single putative causal variant shared between SA and these phenotypes (lifetime smoking index: posterior probability = 0.99, risk-taking behavior: posterior probability = 1) (Table S7 in Supplement 2). Furthermore, a variant in high LD with the chromosome 7 index SNP (rs12666306, $LD r^2 = 0.94$) has a positive genome-wide significant effect on insomnia (reported in GWAS catalog, full summary statistics not available) (Figure 1B; Tables S5 and S6 in Supplement 2). The SA-index SNP has also been implicated in self-harm ideation (52), although not at genome-wide significance, and with a smaller effect size than on SA (Figure 1B).

MAGMA (53) enrichment analyses performed on the primary SA GWAS (see Supplement 1) showed significant enrichment of SA associations in 7 genes (Table S8 in Supplement 2), including *BTN2A1*, which is a brain-expressed gene (54) located within the MHC, that encodes a plasma-membrane protein. There was no enrichment of SA association signal in any of the biological gene sets tested (Table S9 in Supplement 2) or in the set of genes expressed in any of the 54 tissues from the Genotype-Tissue Expression project (Table S10 in Supplement 2). Examining individual genes, a transcriptome-wide association study (see Supplement 1) found 5 genes for which SA risk alleles were significantly associated with brain gene expression: *ERC2*, *RP11-266A24.1*, *TIAF1*, *BACE2*, and *NUFIP2* ($p < 4.28 \times 10^{-6}$) (Table S11 in Supplement 2). None of these genes were within genome-wide significant loci.

### Evidence for Substantial Proportion of SNP-Heritability of SA Not Mediated by Psychiatric Disorders

$h_{SNP}^2$ based on the SA-EUR GWAS was 7.5% (SE = 0.006, $p = 3.02 \times 10^{-40}$) on the liability scale (Table S12 in Supplement 2). Conditioning SA-EUR on MDD resulted in a 45% decrease in the $h_{SNP}^2$ of SA to 4.1% (SE = 0.005, $p = 1.20 \times 10^{-16}$) on the liability scale (Table S12 in Supplement 2). Conditioning on BIP and SCZ in addition to MDD did not further change the $h_{SNP}^2$ estimate ($h_{SNP}^2 = 4.1\%$, SE = 0.005, $p = 1.20 \times 10^{-16}$). The SA-EUR|MDD results showed comparable $h_{SNP}^2$ and complete $r_g$ with a direct GWAS of SA within psychiatric diagnosis
A

<figure><img src="image_3.png" /></figure>

B

<figure><img src="image_4.png" /></figure>

**Figure 1.** Genome-wide significant locus contributes to SA more strongly than psychiatric disorders and other traits. (A) Manhattan plot: the x-axis shows genomic position, and the y-axis shows statistical significance as $-\log_{10}(p$ value). The gray points in the background depict the results of SA-EUR, and the colored points in the foreground depict the results after conditioning these results on MDD (SA-EUR|MDD). The horizontal line shows the genome-wide significance threshold ($p < 5.0 \times 10^{-8}$). (B) Forest plot: the points indicate the log odds ratio of the A allele at rs62474683 (SA-index single nucleotide polymorphism on chromosome 7) on each phenotype, and the error bars show the standard error. The *p* value of association with each phenotype is shown above the error bars. *For insomnia, the effect size of a variant in high linkage disequilibrium with the index single nucleotide polymorphism is shown instead (rs12666306 A allele, linkage disequilibrium $r^2 = 0.94$ with rs62474683 A allele). MDD, major depressive disorder; MHC, major histocompatibility complex; OR, odds ratio; SA, suicide attempt; SA-EUR, European-only suicide attempt meta-analysis; SA-EUR|MDD, SA-EUR results after conditioning on MDD.*

(Supplement 1), confirming the validity of the statistical con-
ditioning approach to control for the genetic effects of psy-
chiatric disorders.

Significant Genetic Overlap Between SA and
Psychiatric Traits or Disorders

Genetic correlations were calculated to explore the genetic
overlap between SA and 12 psychiatric traits or disorders
before and after conditioning on MDD. The SA-EUR GWAS
showed significant r<sub>g</sub> with 11 traits or disorders tested, most
strongly with self-harm ideation (r<sub>g</sub> = 0.82, SE = 0.07, p = 3.57
× 10<sup>-36</sup>), MDD (r<sub>g</sub> = 0.78, SE = 0.04, p = 4.11 × 10<sup>-106</sup>), and
posttraumatic stress disorder (r<sub>g</sub> = 0.74, SE = 0.09, p = 5.29 ×
10<sup>-17</sup>) (Figure 2; Table S13 in Supplement 2). Moderate ge-
netic correlations were also observed between SA and SCZ,
attention-deficit/hyperactivity disorder, BIP, posttraumatic
stress disorder, and alcohol dependence (r<sub>g</sub>s 0.45-0.74)
(Figure 2; Table S13 in Supplement 2).

To investigate whether these genetic correlations were
mediated by MDD, we estimated r<sub>g</sub> with the same traits and

disorders using the SA-EUR|MDD results. Most genetic
correlations with psychiatric disorders remained significant
after conditioning, except for autism spectrum disorder and
Tourette syndrome (Figure 2; Table S13 in Supplement 2). As
expected, the r<sub>g</sub> with MDD significantly decreased after
conditioning (p = 8.4 × 10<sup>-22</sup> block jackknife), as did the r<sub>g</sub>s
with self-harm ideation, posttraumatic stress disorder, and
autism spectrum disorder (Figure 2; Table S13 in
Supplement 2). The remaining psychiatric disorders did
not show Bonferroni corrected significant differences in r<sub>g</sub>
after conditioning on MDD. Because conditional analysis
only removes SNP effects on SA mediated by MDD,
the remaining r<sub>g</sub> between SA-EUR|MDD and MDD (r<sub>g</sub> = 0.53,
SE = 0.06, p = 8.9 × 10<sup>-19</sup>) indicates pleiotropic SNP
effects.

Substantial Shared Genetic Architecture of SA and
Nonpsychiatric Risk Factors Not Mediated by MDD

To assess the shared genetic architecture of SA, psychiatric,
and nonpsychiatric phenotypes, we calculated genetic
<figure><img src="image_5.png" /><figcaption>**Figure 2.** Substantial genetic correlation of SA with psychiatric traits or disorders before and after conditioning SA on MDD. Unfilled points indicate genetic correlations that did not pass the Bonferroni-corrected significance threshold ($p < 4.17 \times 10^{-3}$). Error bars represent the standard error. $p$ values indicate significant differences in genetic correlation after conditioning that pass Bonferroni correction. ADHD, attention-deficit/hyperactivity disorder; AlcUse Disorder P, Alcohol Use Disorders Identification Test-P (measure of problematic consequences of drinking); AN, anorexia nervosa; ASD, autism spectrum disorder; BIP, bipolar disorder; MDD, major depressive disorder; OCD, obsessive-compulsive disorder; PTSD, posttraumatic stress disorder; SA, suicide attempt; SA-EUR, European-only suicide attempt meta-analysis; SA-EUR|MDD, SA-EUR results after conditioning on MDD; SCZ, schizophrenia.</figcaption></figure>

correlations of SA with 768 nonoverlapping phenotypes (46). There were 198 phenotypes that showed a significant $r_g$ with SA-EUR, 133 of which were in one of the predefined SA risk categories (Figure 3A; Table S14 in Supplement 2). The most significant genetic correlations were predominantly with traits related to depressive symptoms, smoking, and socioeconomic status. On examining phenotypes in the risk categories after conditioning on MDD, 110 phenotypes retained a significant $r_g$ with SA-EUR|MDD (Table S14 in Supplement 2). Within the psychiatric risk category, there was a 38% average decrease in the magnitude of genetic correlations with SA-EUR after conditioning, whereas the $r_g$ values in other risk categories were much less affected by conditioning (smoking: 4.6% decrease, education/cognition: 3% decrease, alcohol: 14.5% decrease, and socioeconomic: 9.3% decrease) (Figure 3B).

DISCUSSION

We present a GWAS of SA in over 29,000 cases, identifying 2 genome-wide significant loci, including one more strongly associated with SA than psychiatric disorders or related traits. We demonstrate that a substantial proportion of the SNP-heritability of SA is independent of psychiatric diagnosis. Finally, we show that the genetic liability to SA not mediated by psychiatric disorders is shared with the genetic architecture of nonpsychiatric risk factors.

The locus most strongly associated with SA was in an intergenic region on chromosome 7. The index SNP had a larger effect on SA than on any common psychiatric disorder, remained genome-wide significant after conditioning on MDD, and replicated in an independent cohort from the Million Veteran Program. Taken together, these results

suggest that the genetic association with SA at this locus is not mediated through risk for psychiatric disorders. Functional genomic data do not clearly link this variant to any gene, with the nearest gene being a long noncoding RNA (LINC01392) 149 kb away. The index SNP (rs62474683) is a methylation quantitative trait locus, with the SA risk allele associated with decreased methylation of a nearby DNA methylation site (probe cg04544267) in blood (55). However, this methylation site has not been linked to any gene transcript. Intriguingly, SA risk alleles at this locus have been implicated at genome-wide significance in risk-taking behavior (51), smoking (50), and insomnia (56). While variants in the MHC also reached genome-wide significance for SA, this effect did not remain after conditioning on MDD, suggesting that this association may be a byproduct of psychiatric diagnosis. Indeed, variants in the MHC have previously been associated with risk for a range of psychiatric disorders, including MDD (57).

Our GWAS results provide robust evidence of the $h_{SNP}^2$ of SA, with an estimate of 6.8% on the liability scale (7.5% based on SA-EUR). Importantly, conditioning on MDD resulted in a smaller but significant $h_{SNP}^2$ estimate (4.1%), corroborating previous reports (25,27) of the independent genetic contribution to SA, and illustrating the importance of accounting for potential confounding from the genetics of psychiatric disorders. Traditionally, GWASs have sought to dissect the specific genetic component of SA by studying SA within psychiatric diagnosis or covarying for cases' psychiatric diagnoses (27). Here, we demonstrate that statistical conditioning is an appropriate and easily applicable approach to control for the genetic effects of psychiatric disorders, producing equivalent results to a direct GWAS of SA within psychiatric diagnosis (Supplement 1).
A

<figure><img src="image_6.png" /></figure>

B

<figure><img src="image_7.png" /><figcaption>**Figure 3.** Conditioning SA on MDD reduces genetic correlation with psychiatric phenotypes but has limited effect on other traits. (A) Comparison of significant genetic correlations with the SA-EUR vs. genetic correlations with SA-EUR|MDD. Data include 198 significant genetic correlations after Bonferroni correction (p < .05/768 = 6.51 × 10⁻⁵) annotated by risk category. (B) Top 30 phenotypes with the most significant genetic correlations with SA-EUR before (gray) and after (red) conditioning on MDD (SA-EUR|MDD). Full genetic correlation results, including standard errors, are provided in Table S14 in Supplement 2. GP, general practitioner; MDD, major depressive disorder; SA, suicide attempt; SA-EUR, European-only suicide attempt meta-analysis; SA-EUR|MDD, SA-EUR results after conditioning on major depressive disorder.</figcaption></figure>

SA showed substantial positive genetic correlation with many psychiatric disorders, the highest being with MDD ($r_g$ = 0.78, SE = 0.03), consistent with previous reports (28,29,31). Genetic overlap was also particularly strong with posttraumatic stress disorder, attention-deficit/hyperactivity disorder, SCZ, and BIP ($r_g$ = 0.44 – 0.74). After conditioning on MDD, there was a modest decrease in the genetic correlation of SA with most psychiatric disorders. Notably, SA remained strongly genetically correlated with MDD ($r_g$ = 0.53, SE = 0.06, p = 8.85 × 10⁻¹⁹), representing pleiotropic effects between them. This genetic correlation would only be eliminated if all SNP effects on SA were mediated by MDD. Pleiotropy between psychiatric disorders is widespread (58,59), and accordingly, genetic overlap between SA and related disorders is anticipated. Our findings suggest that many pleiotropic genetic variants increase the risk for SA directly, independent of their effects on psychiatric disorders.

Significant genetic overlap was found between SA and many nonpsychiatric traits, including smoking, lower socioeconomic status, pain, lower educational attainment, reproductive traits, risk-taking behavior, sleep disturbances, and poorer overall general health. While conditioning SA on MDD reduced genetic correlations with psychiatric disorders, the genetic correlation of SA with most nonpsychiatric traits remained unchanged. This suggests a shared genetic architecture between SA and these risk factors that is not mediated by psychiatric illness. There is substantial epidemiological literature on the relationship between sleep disorders (12–15), smoking (18–20), and socioeconomic factors (60–62) and risk for SA but less on genetic overlap between them. We have not examined potential causal relationships between these risk factors and SA, but future

Mendelian randomization studies that will become possible with further increases in the power of SA GWAS may highlight modifiable risk factors.

Several limitations of our study must be noted. Cases were defined using a variety of diagnostic interviews, self-report, or hospital records, which may result in phenotypic heterogeneity. However, suicidal intent was central to all phenotype definitions, and a previous study found 98% concordance between self-report of lifetime SA and face-to-face clinician interview (63). Our GWAS included both nonfatal SA and suicide death cases, and these phenotypes were highly but imperfectly genetically correlated ($r_g$ = 0.77). Genetic correlations between SA and psychiatric disorders were examined using publicly available GWAS summary statistics; however, the prevalence of SA among the cases in these studies is unknown. Finally, population, demographic, and environmental factors are always present in genetic analyses, and while our sample is large and diverse, we did not have sufficient data to assess their possible contribution or confounding effects.

This first collaborative SA GWAS by the ISGC is almost 5-fold larger than previous studies, substantially improving statistical power. We identify a robustly associated SA risk locus and demonstrate genetic liability to SA that is not mediated through psychiatric disorders but is shared with known risk factors. We emphasize that genetic risk does not currently have meaningful predictive utility for SA, and its premature use in clinical or direct-to-consumer settings could be harmful. Future larger studies dissecting the genetic etiology of SA, psychiatric disorders, and other risk factors will provide further insights into the biological mechanisms of risk and assess potential clinical utility.
ACKNOWLEDGMENTS AND DISCLOSURES

Statistical analyses were carried out on the NL Genetic Cluster Computer (http://www.geneticcluster.org) hosted by SURFsara and the Mount Sinai high performance computing cluster (http://hpc.mssm.edu), which is supported by the Office of Research Infrastructure of the National Institutes of Health (Grant Nos. S10OD018522 and S10OD026880). This work was conducted in part using the resources of the Advanced Computing Center for Research and Education at Vanderbilt University, Nashville, TN. This work was funded by the National Institutes of Health (Grant Nos. R01MH116269 and R01MH121455 [to DMR]), NIGMS of the National Institutes of Health (Grant No. T32GM007347 [to JK]), and the Brain & Behavior Research Foundation (NARSAD Young Investigator Award No. 29551 [to NM]).

The content is solely the responsibility of the authors and does not
necessarily represent the official views of any funding body.

We thank the participants who donated their time, life experiences, and DNA to this research, and the clinical and scientific teams that worked with them.

The International Suicide Genetics Consortium has made genome-wide summary results from this study available online (https://tinyurl.com/ISGC2021). This study included some publicly available datasets accessed through dbGaP (Psychiatric Genomics Consortium [PGC] bundle phs001254) and the Haplotype Reference Consortium reference panel v.1.0 (http://www.haplotype-reference-consortium.org/home). Databases used: Open Targets Genetics web portal (https://genetics.opentargets.org), LDHub (http://ldsc.broadinstitute.org), and FUMA (https://fuma.ctglab.nl).

In the past 3 years, RCK was a consultant for Datastat, Inc., Sage Pharmaceuticals, and Takeda. HRK and JG are named as inventors on PCT patent application #15/878,640 entitled: "Genotype-guided dosing of opioid agonists," filed January 24, 2018. HRK is a member of an advisory board for Dicerna Pharmaceuticals and of the American Society of Clinical Psycho-pharmacology's Alcohol Clinical Trials Initiative, which was supported in the last 3 years by AbbVie, Alkermes, Dicerna, Ethypharm, Indivior, Lilly, Lundbeck, Otsuka, Pfizer, Arbor, and Amygdala Neurosciences. DL is an employee of Janssen Research & Development, LLC, and shareholder in Johnson & Johnson, the parent company of the Janssen companies. DL declares that, except for income received from her primary employer, no financial support or compensation has been received from any individual or corporate entity over the past 3 years for research or professional service, and there are no personal financial holdings that could be perceived as constituting a potential conflict of interest. MBS has in the past 3 years been a consultant for Actelion, Acadia Pharmaceuticals, Aptinyx, Bionomics, BioXcel Therapeutics, Epivario, GW Pharmaceuticals, Janssen, Jazz Pharmaceuticals, and Oxeia Biopharmaceuticals. MBS has stock options in Oxeia Biopharmaceuticals and Epivario. HJG has received travel grants and speaker honoraria from Fresenius Medical Care, Neuraxpharm, Servier, and Janssen-Cilag as well as research funding from Fresenius Medical Care. OAA is a consultant for HealthLytx and received speaker's honorarium from Lundbeck and Sunovion. RAP is employed by and holds shares in BioMarin Pharmaceuticals. MCO and MJO are supported by a collaborative research grant from Takeda Pharmaceuticals. That support did not contribute to the work described in this manuscript. EHG has served in the speakers' bureau and the advisory board of Takeda (former Shire do Brasil) Pharmaceutical. JAR-Q was on the speakers' bureau and/or acted as consultant for Eli Lilly, Janssen-Cilag, Novartis, Shire, Takeda, Bial, Shionogui, Lundbeck, Almirall, Braingaze, Sincrolab, Medice, and Rubió in the last 5 years. He also received travel awards (air tickets + hotel) for taking part in psychiatric meetings from Janssen-Cilag, Rubió, Shire, Takeda, Shionogui, Bial, Medice, and Eli Lilly. The Department of Psychiatry chaired by him received unrestricted educational and research support from the following companies in the last 5 years: Eli Lilly, Lundbeck, Janssen-Cilag, Actelion, Shire, Ferrer, Oryzon, Roche, Psious, and Rubió. VR was on the speakers' bureau and/or acted as consultant for Takeda and Rubió in the last 5 years. She also received travel awards (air tickets + hotel) for taking part in psychiatric meetings from Rubió, Shire, Takeda, and Lundbeck. MC was on the speakers' bureau and/or acted as consultant for Janssen-Cilag in the last 5 years. He also received travel awards (air tickets + hotel) for taking part in psychiatric meetings from Janssen-Cilag. All other authors report no biomedical financial interests or potential conflicts of interest.

Full acknowledgments are available in Supplement 1.

ARTICLE INFORMATION

From the Department of Genetics and Genomic Sciences (NM, Dpi, PS, EAS), Department of Neuroscience (PS), and Department of Psychiatry (NM, Dpi, RSK, PS, EAS), Icahn School of Medicine at Mount Sinai; Department of Biostatistics (HG), Departments of Psychiatry and Radiology (JJM), and Department of Psychiatry (HG, IO), Columbia University; Department of Psychiatry (KAH), Weill Cornell Medical College; Department of Psychiatry and Behavioral Sciences (TBB, AHF), State University of New York Downstate Medical Center; Department of Psychiatry (PMr), Columbia University College of Physicians and Surgeons; Columbia University College of Physicians and Surgeons (MMW); and Division of Translational Epidemiology (MMW), New York State Psychiatric Institute, New York, New York; Division of Genetic Medicine (JK, DMR), Department of Medicine, Vanderbilt Genetics Institute, and Department of Biomedical Informatics (DMR) and Department of Psychiatry and Behavioral Sciences (DMR), Vanderbilt University Medical Center, Nashville, Tennessee; Department of Genetics and Computational Biology (AIC, MER) and Genetics and Computational Biology (SEM), QIMR Berghofer Medical Research Institute; School of Biomedical Sciences (AIC, MER), Faculty of Medicine, Institute for Molecular Bioscience (EMB, MT, NRW), Queensland Brain Institute (DM, NRW), and Centre for Children's Health Research (EMB), The University of Queensland; School of Psychology and Counseling (DM), Queensland University of Technology, Brisbane, Queensland, Australia; National Institute for Health Research Maudsley Biomedical Research Centre at South London and Maudsley NHS Foundation Trust (JRIC, GB); Institute of Psychiatry, Psychology and Neuroscience (JT), Department of Psychological Medicine; Institute of Psychology, Psychiatry & Neuroscience (EJSSB); Department of Medical & Molecular Genetics (CML); and Social Genetic and Developmental Psychiatry Centre (JRIC, PMu, RAP, MRy, GB, CML), King's College London; National Institute for Health Research Biomedical Research Centre (JT), King's College London and South London and Maudsley National Health Service Foundation Trust; Division of Psychiatry (NB, AM, JP), University College London; and Genetics (RAP), BioMarin Pharmaceuticals; London, United Kingdom; Department of Psychiatry (ACE, ARD, TBB, AHF, KSK), Virginia Commonwealth University, Richmond, Virginia; Department of Psychiatry (DFLY, JG), Veterans Affairs Connecticut Healthcare Center, West Haven, Connecticut; Division of Human Genetics (DFLY, JG), Department of Psychiatry, Yale University School of Medicine, New Haven, Connecticut; Department of Psychiatry and Behavioral Sciences (AL, AP, EBB), Emory University School of Medicine, Atlanta, Georgia; Department of Psychiatry (ASH, EDB, ETM, HC, ARD) and Biomedical Informatics (HC), University of Utah School of Medicine, Salt Lake City, Utah; Centre for Genomics and Personalized Medicine (AST, ADB, DD), Centre for Integrative Sequencing (AST, PBMo, ADB, DD), Department of Biomedicine (AST, ADB, DD), National Centre for Register-Based Research (ZY, PBMo, EA), Centre for Integrated Register-based Research (PBMo, EA), Psychosis Research Unit (OM), and Lundbeck Foundation Initiative for Integrative Psychiatric Research (AST, ADB, AE, DD), iPSYCH, Aarhus University; and Lundbeck Foundation Initiative for Integrative Psychiatric Research (EA, DMH, OM, MN, TW), Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Aarhus, Denmark; Institute of Epidemiology and Preventive Medicine (M-HS, WJC, P-HK), College of Public Health, National Taiwan University; Department of Psychiatry (WJC, H-CC, S-CL, C-ML, P-HK), National Taiwan University Hospital; and Department of Psychiatry (H-GH), National Taiwan University Hospital and College of Medicine, Taipei, Taiwan; School of Psychology (HJW), Curtin University; and Division of Paediatrics (HJW), The University of Western Australia, Perth, Western Australia, Australia; Department of Psychiatry (HJW, LMT, ZY, CMB), Department of Genetics (ZY), and Department of Nutrition (CMB), University of North Carolina at Chapel Hill, Chapel Hill, North Carolina; Division of Psychiatry (MAd, JDH, AMM) and Institute for Genetics and Molecular Medicine (DPo), University of Edinburgh, Edinburgh, United Kingdom; Department of Psychiatry and Psychotherapy (SA, SRI), Universitätsmedizin Berlin; and Department of Psychiatry (SRO), Universitätsmedizin Berlin, Corporate Member of Freie Universität Berlin, Humboldt-Universität zu Berlin, Berlin Institute of Health, Campus Benjamin Franklin, Berlin, Germany; Department of Psychiatry and Biobehavioral Sciences (MG, MKi), Semel Institute, David Geffen School of Medicine, University of California Los Angeles; David Geffen School of Medicine (MSt), University of California Los Angeles; Department of
Psychiatry and Biobehavioral Sciences (MSt), Semel Institute for Neuroscie-
ence and Human Behavior, University of California Los Angeles; and Jane
and Terry Semel Institute for Neuroscience and Human Behavior (RAO), Los
Angeles; Biostatistics Research Center (SJai), Herbert Wertheim School of
Public Health and Human Longevity Science; Department of Psychiatry
(DLB, JRK, CN), Institute for Genomic Medicine (JRK) and Department of
Psychiatry and School of Public Health (MBS), University of California San
Diego, La Jolla; and Department of Psychiatry (WHK) and Center for
Behavioral Genomics (MTT), Department of Psychiatry, University of Cali-
fornia San Diego, San Diego, California; Department of Psychiatry (AHi),
Yokohama City University Graduate School of Medicine, Yokohama, Japan;
Department of Psychiatry (SO, IO), Kobe University Graduate School of
Medicine, Kobe, Japan; Stanley Center for Psychiatric Research (SRi, PHL,
JWS, CCZ), Stanley Center for Psychiatric Research (KCK), and Program in
Medical and Population Genetics (TE, EAS), Broad Institute, Cambridge,
Massachusetts; Analytical and Translational Genetics Unit (SRi, PHL),
Department of Psychiatry (KCK, JWS), Psychiatric and Neurodevelopmental
Genetics Unit (PHL), and Psychiatric and Neurodevelopmental Genetics Unit
(JWS), Massachusetts General Hospital; Department of Epidemiology (KCK,
CCZ) and Department of Environmental Health (ARo), Harvard TH Chan
School of Public Health; and Department of Health Care Policy (RCK),
Harvard Medical School, Boston, Massachusetts; Population Studies Cen-
ter (EBW), Center for Statistical Genetics and Department of Biostatistics
(MBe, LJS), and Survery Research Center (EBW), Institute for Social
Research, University of Michigan, Ann Arbor, Michigan; BioRealm, LLC
(AWB), Walnut, California; Oregon Research Institute (AWB), Eugene, Ore-
gon; Department of Psychiatry (WHB), Center for Neurobiology and
Behavior, and Department of Psychiatry (DWO, HRK), Perelman School of
Medicine, University of Pennsylvania; Perelman School of Medicine (HH),
University of Pennsylvania; Center for Applied Genomics (XC, YG, HH, DL),
Children's Hospital of Philadelphia; and VISN 4 Mental Illness Research,
Education, and Clinical Center (DWO, HRK), Corporal Michael J. Crescenz
VA Medical Center, Philadelphia, Pennsylvania; Department of Psychoso-
matic Medicine and Psychotherapy (MBh, CS) and Department of Genetic
Epidemiology in Psychiatry (SHW, LZ, FS, JCF, TGS, LS, MRe), Central
Institute of Mental Health, Medical Faculty Mannheim, University of Hei-
delberg, Mannheim, Germany; Center for Eating Disorders at Sheppard Pratt
(HB, SCa); and Department of Psychiatry and Behavioral Sciences (TGS)
and Department of Mental Health (AE), Johns Hopkins University School of
Medicine, Baltimore, Maryland; Center for Neuropsychiatric Research
(WJC), National Health Research Institutes, Miaoli County, Taiwan;
Department of Psychiatry (SCo), University of Minnesota, Minneapolis,
Minnesota; Inserm U955, Institut Mondor de recherches Biomédicales,
Laboratoire, Neuro-Psychiatrie Translationnelle, and Fédération Hospitalo-
Universitaire de Précision Médecine en Addictologie et Psychiatrie (Sjam),
and Faculté de Médecine (MLe), University Paris-Est-Créteil, Créteil; Hôpital
Sainte-Anne (PD, PG), GHU Paris Psychiatrie et Neurosciences; Institute of
Psychiatry and Neuroscience of Paris (PD, PG, NR), INSERM U1266, Uni-
versité de Paris; Department of Psychiatry and Addiction Medicine (FB,
MLe), Assistance Publique Hôpitaux de Paris; Paris Bipolar and TRD Expert
Centres (FB), FondaMental Foundation; UMR-S1144 Team 1: Biomarkers of
relapse and therapeutic response in addiction and mood disorders (FB),
INSERM; Psychiatry (FB), Université Paris Diderot; Institut de Psychiatrie
(BC, M-OK), CNRS GDR 3557; Department of Evaluation, Prevention and
Therapeutic innovation (BC, M-OK), GHU Paris Psychiatrie et Neuroscie-
nces; Team Pathophysiology of psychiatric diseases (BC, M-OK), Uni-
versité de Paris, Institute of Psychiatry and Neuroscience of Paris, INSERM
U1266; and INSERM (MLe), Paris, France; Department of Psychiatry (FF-A,
SJ-M), University Hospital Bellvitge-IDIBELL and CIBEROBN; Department of
Psychiatry (MSA, RB, MC, JAR-Q, MRb, VRi, CS-M, LV-R), Hospital Uni-
versitari Vall d'Hebrón; Department of Genetics, Microbiology & Statistics
(MSA, MRb, CS-M), University of Barcelona; Psychiatric Genetics Unit
(MSA, MC, JAR-Q, MRb, CS-M, LV-R), Group of Psychiatry, Mental Health
and Addiction, Vall d'Hebron Research Institute, Universitat Autònoma de
Barcelona; Department of Psychiatry and Legal Medicine (RB, MC, JAR-Q,
VRi), Universitat Autònoma de Barcelona; and Center for Research in
Environmental Epidemiology (MKo), Barcelona, Spain; Department of Psy-
chiatry and Psychotherapy (MMF), Ludwig-Maximilians-University; Schön
Klinik Roseneck affiliated with the Medical Faculty of the University of

Munich (MMF); Department of Translational Research in Psychiatry (EBB,
BM-M), Max Planck Institute of Psychiatry; Institute of Psychiatric Phe-
nomics and Genomics (MBu, TGS), University Hospital, Ludwig-Max-
imilians-University; Department of Psychiatry (IG), University of Munich; Max
Planck Institute of Psychiatry (MI, SK, SL); and Munich Cluster for Systems
Neurology (BM-M), Munich, Germany; Department of Surgery (SG), Faculty
of Medicine, Department of Psychiatry (ASK, JLK, DBW, SK, JSS, CCZ),
Laboratory Medicine and Pathobiology (CCZ), and Institute of Medical
Sciences (CCZ, ASK, JLK, DBW), University of Toronto; Centre for Addiction
and Mental Health (ASK, JLK, SK, JSS); Department of Paediatric Labora-
tory Medicine (CRM) and Department of Genetics and Genomic Biology
(SWS), The Hospital for Sick Children; Centre for Mental Health (DBW) and
Program for Eating Disorders (DBW), University Health Network; Molecular
Brain Science (JBV), Centre for Addiction and Mental Health; and Molecular
Brain Science (CCZ), Campbell Family Mental Health Research Institute,
Centre for Addiction and Mental Health, Toronto, Ontario, Canada;
Department of Psychiatry and Behavioral Sciences (SJG), SUNY Upstate
Medical University, Syracuse, New York; Eating Recovery Center (CJ),
Denver, Colorado; Department of Psychology (PKK), Florida State Univer-
sity, Tallahassee, Florida; Department of Psychology (KLK), Michigan State
University, Lansing, Michigan; Department of Psychiatry and Psychotherapy
(KL), University Medical Center, Mainz, Germany; Department of Clinical
Psychology (LL), The Chicago School of Professional Psychology, Wash-
ington, DC; Behavioral Health Services (SIS), Kaiser Permanente Washing-
ton, Seattle, Washington, DC; BESE Division (PJM), King Abdullah University
of Science and Technology, Thuwal, Saudi Arabia; Department of Psychiatry
(PJM, EC, GP, MP), Lausanne University Hospital and University of Lau-
sanne, Lausanne, Switzerland; Department of Psychiatry and Behavioral
Science (JEM), University of North Dakota School of Medicine and Health
Sciences, Fargo, North Dakota; HudsonAlpha Institute for Biotechnology
(RMM), Huntsville, Alabama; Department of Psychology (VRo), St. Peters-
burg State University; and Department of Borderline Disorders and Psy-
chotherapy (VRo), V.M. Bekhterev National Medical Research Center for
Psychiatry and Neurology; Saint Petersburg, Russian Federation; National
Centre for Suicide Research and Prevention of Mental Ill-Health (MSo, DW),
LIME, Karolinska Institutet; Department of Clinical Neuroscience (IA), Centre
for Psychiatry Research, Karolinska Institutet; Department of Clinical
Neuroscience (LA), Institut of Environmental Medicine (LA), and Department
of Medical Epidemiology and Biostatistics (MLa, CMB), Karolinska Institutet,
Stockholm, Sweden; Department of Clinical Sciences (RA), Psychiatry,
Umeå University Medical Faculty, Umeå, Sweden; Department of Psychi-
atrie Research (IA), Diakonhemmet Hospital; NORMENT (IA), Institute of
Clinical Medicine, University of Oslo; Division of Mental Health and Addic-
tion (OAA, IM, OBS) and Department of Medical Genetics (SD), Oslo Uni-
versity Hospital; NORMENT (OAA, OBS), University of Oslo; and Division of
Mental Health and Addiction (IM), University of Oslo, Institute of Clinical
Medicine, Oslo, Norway; Discipline of Psychiatry (TMA), University of Ade-
laide, Adelaide, South Australia, Australia; Department of Psychiatry (MAI),
Dalhousie University, Halifax, Nova Scotia, Canada; National Institute of
Mental Health (MAI), Klecany, Czech Republic; Psychiatry (AA), Berkshire
Healthcare NHS Foundation Trust, Bracknell, United Kingdom; Institute of
Biological Psychiatry (VA, TW), Copenhagen Mental Health Services,
Copenhagen University Hospital; Lundbeck Foundation Initiative for inte-
grative Psychiatric Research (VA, PBMo), iPSYCH; Danish Research Insti-
tute for Suicide Prevention (AE), Mental Health Centre Copenhagen; Center
for Neonatal Screening (DMH), Department for Congenital Disorders,
Statens Serum Institut; Mental Health Center Copenhagen (MN), Copen-
hagen University Hospital; and Department of Clinical Medicine (TW), and
Lundbeck Foundation GeoGenetics Centre (TW), GLOBE Institute, Univer-
sity of Copenhagen, Copenhagen, Denmark; Biomedical Network Research
Centre on Mental Health (MSA, RB, MC, JAR-Q, MRb, VRi, CS-M), Instituto
de Salud Carlos III, Madrid, Spain; Department of Psychiatry and Psycho-
therapy (SVA, HJG) and Institute for Community Medicine (HV), University
Medicine Greifswald, Greifswald, Mecklenburg-Vorpommern, Germany;
Department of Psychiatry (MHA), University of Coimbra, Coimbra, Portugal;
Laboratory of Developmental Psychiatry (CHDB) and ADHD Outpatient
Program (EHG), Adult Division, Hospital de Clínicas de Porto Alegre;
Department of Genetics (CHDB) and Department of Psychiatry (EHG), Uni-
versidade Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul,
Brazil; Department of Psychiatry (BTB), Melbourne Medical School, Uni-
versity of Melbourne, Melbourne, Victoria, Australia; Department of Psy-
chiatry (BTB) and Institute of Epidemiology and Social Medicine (KB),
University of Münster, Münster, Nordrhein-Westfalen, Germany; Health
Sciences Research (JMB) and Department of Psychiatry & Psychology (MF),
Mayo Clinic, Rochester, Minnesota; Department of Psychiatry (MPB, WC,
RSK), UMC Utrecht Hersencentrum, Utrecht, the Netherlands; School of
Psychology (RB, JMG), School of Psychiatry (MJG, PBMi, GR, CSW, TWW),
and School of Medical Sciences (JMF, PRS), University of New South
Wales; and Neuroscience Research Australia (JMF, JMG, MJG, PRS, CSW,
TWW), Sydney, New South Wales, Australia; Mental Health Unit (JAC),
Department of Psychiatry, Faculty of Medicine, Granada University Hospital
Complex, and Department of Psychiatry (BG), Department of Biochemistry
and Molecular Biology II and Institute of Neurosciences (MRv), and
Department of Nursing (EM), Faculty of Medicine and Biomedical Research
Centre, University of Granada, Granada, Spain; Institute of Neuroscience
and Medicine (SCI, AJF), Research Centre Jülich, Jülich, Germany; Institute
of Medical Genetics and Pathology (SCI, PH) and Department of Biomed-
cine (SCI, PH), University Hospital Basel, Basel, Switzerland; Institute of
Human Genetics (SCI, FD, AJF, SH-H, PH, MMN), University of Bonn,
School of Medicine & University Hospital Bonn, Bonn, Germany; Neuro-
psychiatric Genetics Research Group (AC), Department of Psychiatry and
Trinity Translational Medicine Institute, Trinity College Dublin, Dublin,
Ireland; Medical Research Council for Neuropsychiatric Genetics and
Genomics (NC, MLH, IJ, MCO, MJO, JTRW), Division of Psychological
Medicine and Clinical Neurosciences, Cardiff University, Cardiff, United
Kingdom; Department of Translational Genomics (DC), University of
Southern California, Pasadena, California; NORMENT (SD), KG Jebsen
Centre for Psychosis Research, Department of Clinical Science, University
of Bergen, Bergen, Norway; Department of Medical & Molecular Genetics
(HJE, Indiana University; and Biochemistry and Molecular Biology (HJE),
Indiana University School of Medicine, Indianapolis, Indiana; Centre for
Human Genetics (AJF), University of Marburg, Marburg, Germany; Depart-
ment of Psychiatry and Behavioral Sciences (PVG, ARS), NorthShore Uni-
versity HealthSystem, Evanston, Illinois; Department of Psychiatry and
Behavioral Neuroscience (PVG, ARS), University of Chicago, Chicago, Illi-
nois; Department of Psychiatry, Psychotherapy and Psychosomatics (IG,
AHA, BK, DR), Martin-Luther-University Halle-Wittenberg, Halle, Germany;
Biometric Psychiatric Genetics Research Unit (MG-S), Alexandru Obregia
Clinical Psychiatric Hospital, Bucharest, Romania; Mental Health Depart-
ment (JG-P, FM), University Regional Hospital, Biomedicine Institute, Må-
laga, Spain; Psychiatry (SPH), Kaiser Permanente Northern California, San
Francisco, California; Department of Psychiatry (JH), Laboratory of Psychi-
iatric Genetics, Poznan University of Medical Sciences, Poznań, Poland;
Department of Psychological Medicine (LAJ), University of Worcester,
Worcester, United Kingdom; Department of Psychiatry and Neuroscience
(LJ, MLa), University of Gothenburg, Gothenburg, Sweden; Psychiatry (JLa),
North East London NHS Foundation Trust, Ilford, Cheshire, United Kingdom;
Department of Psychiatry and Behavioral Sciences (DFLn, LMW), Stanford
University, Stanford, California; Department of Human Genetics (CL),
Department of Psychiatry (GT), and Department of Neurology and Neuro-
surgery (GAR), Faculty of Medicine, McGill University; and Montreal
Neurological Institute and Hospital (CL, GAR), Montreal, Québec, Canada;
Cancer Epidemiology and Prevention (JLi). Sklodowska-Curie Cancer
Center and Institute of Oncology, Warsaw, Poland; Research Institute (SLM),
Lindner Center of HOPE, Mason, Ohio; Department of Psychiatry (YM,
BWJHP), Amsterdam UMC, Vrije Universiteit and GGZ in Geest, Amsterdam,
Netherlands; Mental Health (GM), Faculty of Medicine and Health Sciences,
Norwegian University of Science and Technology, NTNU; and Psychiatry
(GM), St. Olavs University Hospital, Trondheim, Norway; University of Liv-
erpool (BM-M), Liverpool, United Kingdom; Psychiatry and Human Genetics
(VN), University of Pittsburgh, Pittsburgh, Pennsylvania; Psychiatry (RAO),
Erasmus University Medical Center, Rotterdam, the Netherlands; College of
Medicine Institute for Genomic Health (CP) and Institute for Genomic Health
(CP, MTP), SUNY Downstate Medical Center College of Medicine, Brooklyn,
New York; Department of Psychiatry (JBP, VW), University of Iowa, Iowa
city, Iowa; Department of Psychiatry (DQ), University of Oxford; and Uni-
versity of Oxford (RAP), St. Edmund Hall, Oxford, United Kingdom;
Department of Psychiatry (ARe), Psychosomatic Medicine and

Psychotherapy, University Hospital Frankfurt, Frankfurt, Germany; Depart-
ment of Physiology and Biophysics (DLR), Instituto de Ciencias Biomedicas
Universidade de Sao Paulo, Sao Paulo, Sao Paulo, Brazil; Human Genetics
Branch (TGS), Intramural Research Program, National Institute of Mental
Health; Department of Psychiatry (RJU), Uniformed University of the Mental
Sciences; and Division of Cancer Epidemiology and Genetics (JS), National
Cancer Institute, Bethesda, Maryland; Department of Psychiatry and Psy-
chotherapy (TGS), University Medical Center Göttingen, Göttingen, Ger-
many; Department of Biomedical and NeuroMotor Sciences (ASe),
University of Bologna, Bologna, Italy; Menninger Department of Psychiatry
and Behavioral Sciences (GS), Baylor College of Medicine, Houston, Texas;
Laboratory of Neuropsychiatry (GS), IRCCS Santa Lucia Foundation, Rome,
Italy; Department of Environmental Epidemiology (BS), Nofer Institute of
Occupational Medicine, Lódz, Poland; Duke Molecular Physiology Institute
(AEA-K, ERH, MAH, XQ) and Department of Psychiatry and Behavioral
Sciences (JCB, NAK), Duke University Medical Center; VISN 6 Mid-Atlantic
Mental Illness Research (JCB, NAK) and Cooperative Studies Program
Epidemiology Center, Education, and Clinical Center (ERH), and VA Health
Services Research and Development Center of Innovation to Accelerate
Discovery and Practice Transformation (JHL), Durham Veterans Affairs
Health Care System, Durham, North Carolina; Theoretical Division (BM), Los
Alamos National Laboratory, Los Alamos National Laboratory, Los Alamos,
New Mexico; Center of Mental Health Research (AE), Australian National
University, Canberra, Australia; Estonian Genome Center (TE), Institute of
Genomics, University of Tartu, Tartu, Estonia; Neuroscience (QSL), Janssen
Research & Development, LLC, Titusville, New Jersey; Department of Ge-
netics and Computational Biology (NGM) and Department of Population
Health (CMO, DCW), QIMR Berghofer Medical Research Institute, Herston,
Brisbane, Queensland, Australia; and Institute of Health and Wellbeing
(DJS), University of Glasgow, Glasgow, United Kingdom.

Major Depressive Disorder Working Group of the Psychiatric Ge-
nomics Consortium

Naomi R. Wray*¹,², Stephan Ripke*³,⁴,⁵, Manuel Mattheisen*⁶,⁷,⁸, Maciej Trzaskowski¹, Enda M. Byrne¹, Abdel Abdellaoui⁹, Mark J. Adams¹⁰, Esben Agerbo¹¹,¹²,¹³, Tracy M. Air¹⁴, Till F.M. Andlauer¹⁵,¹⁶, Silviu-Alin Bacanu¹⁷, Marie Bækvad-Hansen¹³,¹⁸, Aartjan T.F. Beekman¹⁹, Tim B. Bigdeli¹⁷,²⁰, Elisabeta B. Binder¹⁵,²¹, Julien Bryois²², Henriette N. Buttenschön¹³,²³,²⁴, Jonas Bybjerg-Grauholm¹³,¹⁸, Na Cai²⁵,²⁶, Enrique Castelao²⁷, Jane Hvarregaard Christensen⁸,¹³,²⁴, Toni-Kim Clarke¹⁰, Jonathan R.I. Coleman²⁸, Lucia Colodro-Conde²⁹, Baptiste Couvy-Duchesne²,³⁰, Nick Craddock³¹, Gregory E. Crawford³²,³³, Gail Davies³⁴, Franziska Degenhardt³⁵, Eske M. Derks²⁶, Nese Direk³⁶,³⁷, Conor V. Dolan⁹, Erin C. Dunn³⁸,³⁹,⁴⁰, Thalia C. Eley²⁸, Valentina Escott-Price⁴¹, Farnush Farhadi Hassan Kiadeh⁴², Hilary K. Finucane⁴³,⁴⁴, Jerome C. Foo⁴⁵, Andreas J. Forstner³⁵,⁴⁶,⁴⁷,⁴⁸, Josef Frank⁴⁵, Héléna A. Gaspar²⁸, Michael Gill⁴⁹, Fernando S. Goes⁵⁰, Scott D. Gordon²⁹, Shantel Marie Weinsheimer¹³,⁵⁴, Jürgen Wellmann¹⁰¹, Gonneke Willemsen⁹, Stephanie H. Witt⁴⁵, Yang Wu¹, Hualin S. Xi¹¹², Jian Yang²,¹¹³, Futao Zhang¹, Volker Arolt¹¹⁴, Bernhard T. Baune¹¹⁴,¹¹⁵,¹¹⁶, Klaus Berger¹⁰¹, Dorret I. Boomsma⁹, Sven Cichon³⁵,⁴⁷,¹¹⁷,¹¹⁸, Udo Dannlowski¹¹⁴, E.J.C. de Geus⁹,¹¹⁹, J. Raymond Depaulo⁵⁰, Enrico Domenici¹²⁰, Katharina Domschke¹²¹,¹²², Tónu Esko⁵,⁷⁸, Hans J. Grabe¹⁰⁹, Steven P. Hamilton¹²³, Jakob Grove⁸,¹³,²⁴,⁵¹, Lynsey S. Hall¹⁰,⁵², Christine Sohlem Hansen¹³,¹⁸, Thomas F. Hansen⁵³,⁵⁴,⁵⁵, Stefan Herms³⁵,⁴⁷, Ian B. Hickie⁵⁶, Per Hoffmann³⁵,⁴⁷, Georg Homuth⁵⁷, Carsten Horn⁵⁸, Jouke-Jan Hottenga⁹, David M. Hougaard¹³,¹⁸, David M. Howard¹⁰,²⁶, Marcus Isling⁵⁹, Rick Jansen¹⁹, Ian Jones⁶⁰, Lisa A. Jones⁶¹, Eric Jorgenson⁶², James A. Knowles⁶³, Isaac S. Kohane⁶⁴,⁶⁵,⁶⁶, Julia Kraft⁴, Warren W. Kretzschmar⁶⁷, Zoltán Kutalik⁶⁸,⁶⁹, Yihan Li⁶⁷, Penelope A. Lind²⁹, Donald J. MacIntyre⁷⁰,⁷¹, Dean F. Mackinnon⁵⁰, Robert M. Maier², Wolfgang Maier⁷², Jonathan Marchini⁷³, Hamdi Mbarek⁹, Patrick McGrath⁷⁴, Peter McGuffin²⁸, Sarah E. Medland²⁹, Divya Mehta²,⁷⁵, Christel M. Middeldorp⁹,⁷⁶,⁷⁷, Evelin Mihailov⁷⁸, Yuri Milaneschi¹⁹, Lili Milani⁷⁸, Francis M. Mondimore⁵⁰, Grant W. Montgomery¹, Sara Mostafavi⁷⁹, ⁸⁰, Niamh Mullins²⁸, Matthias Nauck⁸¹,⁸², Bernard Ng⁸⁰, Michel G. Nivard⁹, Dale R. Nyholt⁸³, Paul F. O'Reilly²⁸, Hogni Oskarsson⁸⁴, Caroline Hayward¹²⁴, Andrew C. Heath⁸⁹, Kenneth S. Kendler¹⁷, Stefan Kloiber⁵⁹,¹²⁵,¹²⁶, Glyn Lewis¹²⁷, Qingqin S. Li¹²⁸, Susanne Lucae⁵⁹, Pamela A.F. Madden⁸⁹, Patrik K. Magnusson²², Nicholas G. Martin²⁹, Andrew M. McIntosh¹⁰,³⁴, Andres Metspalu⁷⁸,¹²⁹, Ole Mors¹³,¹³⁰, Preben Bo Mortensen¹¹,¹²,¹³,¹²⁴, Bertram Müller-Myhsok¹⁵,¹³¹,¹³², Merete Nordentoft¹³,¹³³,
Markus M. Nöthen³⁵, Michael C. O'Donovan⁶⁰, Sara A. Paciga¹³⁴, Nancy L. Pedersen²², Michael J. Owen⁶⁰, Jodie N. Painter²⁹, Carsten Bøcker Pedersen¹¹,¹²,¹³, Marianne Ørgtj Pedersen¹¹,¹²,¹³, Roseann E. Peterson¹⁷,⁸⁵, Wouter J. Peyrot¹⁹, Giorgio Pisticz²⁷, Danielle Posthuma⁸⁶,⁸⁷, Jorge A. Quiroz⁸⁸, Per Qvist⁸,¹³,²⁴, John P. Rice⁸⁹, Brien P. Riley¹⁷, Margarita Rivera²⁸,⁹⁰, Saira Saeed Mirza³⁶, Robert Schoevers⁹¹, Eva C. Schulte⁹²,⁹³, Ling Shen⁶², Jianxin Shi⁹⁴, Stanley I. Shyn⁹⁵, Engilbert Sigurdsson⁹⁶, Grant C.B. Sinnamon⁹⁷, Johannes H. Smit¹⁹, Daniel J. Smith⁹⁸, Hreinn Stefansson⁹⁹, Stacy Steinberg⁹⁹, Fabian Streit⁴⁵, Jana Strohmaier⁴⁵, Katherine E. Tansey¹⁰⁰, Henning Teismann¹⁰¹, Alexander Teumer¹⁰², Wesley Thompson¹³,⁵⁴,¹⁰³,¹⁰⁴, Pippa A. Thomson¹⁰⁵, Thorgeir E. Thorgeirsson⁹⁹, Matthew Traylor¹⁰⁶, Jens Treutlein⁴⁵, Vassily Trubetskoy⁴, André G. Uitterlinden¹⁰⁷, Daniel Umbricht¹⁰⁸, Sandra Van der Auwera¹⁰⁹, Albert M. van Hemert¹¹⁰, Alexander Viktorin²², Peter M. Visscher¹,², Yunpeng Wang¹³,⁵⁴,¹⁰⁴, Bradley T. Webb¹¹¹, Brenda W.J.H. Penninx¹⁹, Roy H. Perlis³⁸,¹³⁵, David J. Porteous¹⁰⁵, James B. Potash¹³⁶, Martin Preisig²⁷, Marcella Rietschel⁴⁵, Catherine Schaefer⁶², Thomas G. Schulze⁴⁵,⁹³,¹³⁷,¹³⁸,¹³⁹, Jordan W. Smoller³⁸,⁹⁹,⁴⁰, Kari Stefansson⁹⁹,¹⁴⁰, Henning Tiemeier³⁶,¹⁴¹,¹⁴², Rudolf Uher¹⁴³, Henry Völzke¹⁰², Myrna M. Weissman⁷⁴,¹⁴⁴, Thomas Werge¹³,⁵⁴,¹⁴⁵, Cathryn M. Lewis²⁸,¹⁴⁶, Douglas F. Levinson*¹⁴⁷, Gerome Breen*²,⁸⁴,¹⁸⁴, Anders D. Bøglum*⁸,¹³,²⁴, and Patrick F. Sullivan*²²,¹⁴⁹,¹⁵⁰

See Supplement 1 for all affiliations of consortium members.

## Bipolar Disorder Working Group of the Psychiatric Genomics Consortium

Niamh Mullins¹,², Andreas J. Forstner³,⁴,⁵, Kevin S. O'Connell⁶,7, Branton Coombes⁸, Jonathan R.I. Coleman⁹,¹⁰, Zhen Qiao¹¹, Thomas D. Als¹²,¹³,¹⁴, Tim B. Bigdeli¹⁵,¹⁶, Sigrid Borte¹⁷,¹⁸,¹⁹, Julien Bryois²⁰, Alexander W. Charney², Ole Kristian Drange²¹,²², Michael J. Gandal²³, Saskia P. Hagenaars⁹,¹⁰, Masashi Ikeda²⁴, Nolan Kamitaki²⁵,²⁶, Minsoo Kim²³, Kristi Krebs²⁷, Georgia Panagiotaropoulou²⁸, Brian M. Schilder¹,²⁹,³⁰,³¹, Laura G. Sloofman¹, Stacy Steinberg²³, Vassily Trubetskoy²⁸, Bendik S. Winsvold¹⁹,³³, Hong-Hee Won³⁴, Liliya Abramova³⁵, Kristina Adorjan³⁶,³⁷, Esben Agerbo⁴⁴,¹³,⁸⁹,³⁹, Mariam Al Eissa⁴⁰, Diego Albani⁴¹, Ney Alliey-Rodriguez⁴²,⁴³, Adebayo Anjorin⁴⁴, Verner Antilla⁴⁵, Anastasia Antoniou⁴⁶, Swapail Awasthi²⁸, Ji Hyun Baek⁴⁷, Marie Bækvad-Hansen¹⁴,⁸⁸, Nicholas Bass⁴⁰, Michael Bauer⁴⁹, Eva C. Beins⁵¹, Sarah E. Bergen²⁰, Armin Birner²⁰, Carsten Bøcker Pedersen¹⁴,¹³,⁸⁹,³⁹, Erlend Boen⁵¹, Marco P. Boks⁵², Rosa Bensch³⁵,⁵⁴,⁵⁵,⁵⁶, Murielle Brum⁵⁷, Ben M. Brumenton¹⁹, Nathalie Brunkhorst-Kanaan⁵⁷, Monika Budde³⁶, Jonas Bybjerg-Grauholm¹⁴,⁸⁸, William Byerley⁵⁸, Murray Cairns⁵⁹, Miquel Casas⁵³,⁵⁴,⁵⁵,⁵⁶, Pablo Cervantes⁶⁰, Toni-Kim Clarke⁶¹, Cristiana Cruceanu⁶⁰,⁶², Alfredo Cuellar-Barboza⁶³,⁶⁴, Julie Cunningham⁶⁵, David Curtis⁶⁶,⁶⁷, Piotr M. Czerski⁶⁸, Anders M. Dale⁷⁰, Nina Dalkner⁵⁰, Friederike S. David³, Franziska Degenhardt³,⁷⁰, Srdjan Djurovic¹⁷,¹⁷, Amanda L. Dobbyn¹,², Athanassios Douzenis⁴⁶, Torbjørn Elvsåshagen¹⁸,⁷³,⁷⁴, Valentina Escott-Price⁷⁵, I. Nicol Ferrier⁷⁶, Alessia Fiorino⁴⁰, Tatiana M. Foroud⁷⁷, Liz Forty⁷⁸, Oleksandr Frei⁶,⁸¹, Nelson B. Freimer²³,⁷⁹, Louise Frisén⁸⁰, Katrin Gade³⁶,⁸¹, Julie Garnham⁸², Joel Gelernter⁸³,⁸⁴,⁸⁵, Marianne Giortz Pedersen¹⁴,¹³,⁸⁹,³⁹, Ian R. Gizer⁸⁶, Scott D. Gordon⁸⁷, Katherine Gordon-Smith⁸⁸, Tiffany A. Greenwood⁸⁹, Jakob Grove¹²,¹³,¹⁴,⁹⁰, José Guzman-Parra⁹¹, Kyooseob Ha⁹², Magnus Haldsson⁹³, Martin Hautzinger⁹⁴, Urs Heilbronner⁹⁵, Dennis Hellgren²⁰, Stefan Herms⁹,⁵⁹,⁶⁰, Per Hoffmann⁹,⁵⁹,⁶⁰, Peter A. Holms⁷⁵, Laura Hucks¹,², Stéphane Jamain⁹⁷,⁹⁸, Jessica S. Johnson¹,², Janos L. Kalman³⁶,³⁷,⁹⁹, Yochi Kamatani¹⁰⁰,¹⁰¹, James L. Kennedy¹⁰²,¹⁰³,¹⁰⁴,¹⁰⁵, Sarah Kittel-Schneider⁵⁷,¹⁰⁶, James A. Knowles¹⁰⁷,¹⁰⁸, Manolis Kogevinas¹⁰⁹, Maria Koromina¹¹⁰, Thorsten M. Kranz⁵⁷, Henry R. Kranzler¹¹¹,¹¹¹, Michiaki Kubo¹¹³, Ralph Kupka¹¹⁴,¹¹⁵,¹¹⁶, Steven A. Kushner¹¹⁷, Catharina Lavibratt¹¹⁸,¹¹⁹, Jacob Lawrence¹²⁰, Markus Leber¹²¹, Heon-Jeong Lee¹²², Phil H. Lee¹²³, Shawn E. Levy¹²⁴, Catrin Lewis⁷⁵, Calwing Liao¹²⁵,¹²⁶, Susanne Lucae⁶², Martin Lundberg¹¹⁸,¹¹⁹, Donald J. Maclntyre¹²⁷, Sigurdur H. Magnusson³², Wolfgang Maier¹²⁸, Adam Maihofer⁸⁹, Dolores Malaspinca¹,², Eirini Maratou¹²⁹, Lina Martinsson⁸⁰, Manuel Mattheisen¹²,¹³,¹⁴,¹⁰⁶,¹³⁰, Nathaniel W. McGregor¹³¹, Peter McGuffin⁹, James D. McKay¹³², Helena Medeiros¹⁰⁸, Sarah E Medland⁸⁷, Vincent Millischer¹¹⁸,¹¹⁹, Grant W. Montgomery¹¹, Jennifer L. Moran²⁵,¹³³, Derek W. Morris¹³⁴, Thomas W. Mühleisen⁴,⁵⁹, Niamh O'Brien⁴⁰, Claire O'Donovan⁸², Loes M. Olde Loohuis²³,⁷⁹, Lilijana Oruc¹³⁵, Sergi Papiol³⁶,³⁷, Antonio F. Díazinas⁷⁵, Amy Perry⁸⁸, Andrea Pfennig⁴⁹, Evgenia Porichi⁴⁶, James B. Potash¹³⁶, Digby Quested¹³⁷,¹³⁸, Towfique Raj¹,²⁹,³⁰,³¹, Mark H. Rapaport¹³⁹, J. Raymond DePaulo¹³⁶, Eline J.

Regeer¹⁴⁰, John P. Rice¹⁴¹, Fabio Rivas⁹¹, Margarita Rivera¹⁴²,¹⁴³, Julian Roth¹⁰⁶, Panos Roussos¹,²,²⁹, Douglas M. Ruderfer¹⁴⁴, Cristina Sánchez-Mora⁵³,⁵⁴,⁵⁶,¹⁴⁵, Eva C. Schulte³⁶,³⁷, Fanny Senner³⁶,³⁷, Sally Sharp⁴⁰, Paul D. Shilling⁸⁹, Engilbert Sigurdsson⁹³,¹⁴⁶, Lea Sigrignano⁷⁸, Claire Slaney⁸², Olav B. Smeland⁶,⁷, Daniel J. Smith¹⁴⁷, Janet L. Sobell¹⁴⁸, Christine Søholm Hansen¹⁴,⁸⁸, Maria Soler Artigas⁵³,⁵⁴,⁵⁶,¹⁴⁵, Anne T. Spijker¹⁴⁹, Dan J. Stein¹⁵⁰, John S. Strauss¹⁰², Beata Świątkowska¹⁵¹, Chikashi Terao¹⁰¹, Thorgeir E. Thorgeirsson³², Claudio Toma¹⁵²,¹⁵³,¹⁵⁴, Paul Tooney⁵⁹, Evangelia-Eirini Tsermpini¹¹⁰, Marquis P. Vawter¹⁵⁵, Helmut Vedder¹⁵⁶, James T.R. Walters⁷⁵, Stephanie H. Witt⁷⁸, Simon Xi¹⁵⁷, Wei Xu¹⁵⁸, Jessica Mei Kay Yang⁷⁵, Allan H. Young¹⁵⁹,¹⁶⁰, Hannah Young¹, Peter P. Zandi¹³⁶, Hang Zhou⁸³,⁸⁴, Lea Zillich⁷⁸, HUNT All-In Psychiatry¹⁶¹, Rolf Adolfsson¹⁶², Ingrid Agartz⁵¹,¹³⁰,¹⁶³, Martin Alda⁸²,¹⁶⁴, Lars Alfredsson¹⁶⁵, Gulja Badajova¹⁶⁶, Lena Backlund¹¹⁸,¹¹⁹, Bernhard T. Baune¹⁶⁷,¹⁶⁸,¹⁶⁹, Frank Bellivier¹⁷,¹⁷¹, Susanne Bengesser⁵⁰, Wade H. Berretti¹⁷², Douglas H.R. Blackwood⁶¹, Michael Boehnke¹⁷³, Anders D. Bøglum¹⁴,¹⁷⁴,¹⁷⁵, Gerome Breen⁹,¹⁰, Vaughan J. Carr¹⁷⁶, Stanley Catts¹⁷⁷, Aiden Corvin¹⁷⁸, Nicholas Craddock⁷⁵, Udo Dännlowski¹⁶⁷, Dimitris Dikeos¹⁷⁹, Tõnu Esko²⁶,²⁷,¹⁸⁰,¹⁸¹, Aruno Etain¹⁷⁰,¹⁷¹, Panagiotis Ferentinos⁹,⁴⁶, Mark Fraye⁶⁴, Janice M. Fullertón¹⁵²,¹⁵³, Micha Gawlik¹⁰⁶, Elliot S. Gershon⁴²,¹⁸², Fernando S. Goes¹³⁶, Melissa J. Green¹⁵²,¹⁷⁶, Maria Grigoroiu-Serbanescu¹⁸³, Joanna Hauser⁶⁸, Frans Henskens⁵⁹, Jan Hillert⁸⁰, Kyung Sue Hong⁴⁷, David M. Hougaard¹⁴,⁸⁸, Christina M. Hultman²⁰, Kristian Hveem¹⁹,¹⁸⁴, Nakao Iwata²⁴, Assen V. Jablensky¹⁸⁵, Ian Jones⁷⁵, Lisa A Jones⁸⁸, René S. Kahn²,⁵², John R. Kelsoe⁸⁹, George Kirov⁷⁵, Mikael Landén²⁰,¹⁸⁶, Marion Leboyer⁹⁷,⁹⁸,¹⁸⁷, Cathryn M. Lewis⁹,¹⁰,¹⁸⁸, Qingqin S. Li¹⁸⁹, Jolanta Lissowska¹⁹⁰, Christine Lochner¹⁹¹, Carmel Loughland⁵⁹, Nicholas G. Martin⁸⁷,¹⁹², Carol A. Mathews¹⁹³, Fermin Mayoral⁹¹, Susan L. McElroy¹⁹⁴, Andrew M. McIntosh¹²⁷,¹⁹⁵, Francis J. McMahon¹⁹⁶, Ingrid Melle⁶,¹⁹⁷, Patricia Michie⁵⁹, Lili Milani²⁷, Philip B. Mitchell¹⁷⁶, Gunnar Morken²¹,¹⁹⁸, Ole Mors¹⁴,¹⁹⁹, Preben Bo Morgenstern¹²,¹⁴,¹³,⁸⁹,³⁹, Bryan Mowry¹⁷⁷, Bertram Müller-Myhks²⁶,²⁰⁰,²⁰¹, Richard M. Myers¹²⁴, Benjamin M. Neale²⁵,⁴⁵,¹⁸⁰, Caroline M. Nievergelt⁸⁹,²⁰², Merete Nordentoft¹⁴,²⁰³, Markus M. Nöthen³, Michael C. O'Donovan⁷⁵, Ketil J. Oedegaard²⁰⁴,²⁰⁵, Tomas Olsson²⁰⁶, Michael J. Owen²⁵, Sara A. Paciga²⁰⁷, Chris Pantelis²⁰⁸, Carlos Pato¹⁰⁸, Michele T. Pato¹⁰⁸, George P. Patrinos¹¹⁰,¹⁹⁰,²¹⁰, Roy H. Perlis²¹¹,²¹², Danielle Posthuma²¹³,²¹⁴, Josep Antoni Ramos-Quiroga⁵³,⁵⁴,⁵⁵,⁵⁶, Andreas Reif⁵⁷, Eva Z. Reininghaus⁵⁰, Marta Ribas⁵⁰,⁵³,⁵⁴,⁵⁶,¹⁴⁵, Marcella Rietschel⁷⁸, Stephan Rekke²⁵,²⁸,⁴⁵, Guy A. Rouleau¹²⁶,²¹⁵, Takeo Saito²⁴, Ulrich Schall⁵⁹, Martin Schalling¹¹⁸,¹¹⁹, Peter R. Schofield¹⁵²,¹⁵³, Thomas G. Schulze³⁶,⁷⁸,⁸¹,¹³⁶,²¹⁶, Laura J. Scott¹⁷³, Rodney J. Scott⁵⁹, Alessandro Serretti²¹⁷, Cynthia Shannon Weickert¹⁵²,¹⁷⁶,²¹⁸, Jordan W. Smoller²⁵,¹³³,²¹⁹, Hreinn Stefansson³², Kari Stefansson³²,²²⁰, Eystein Sordal²²¹,²²², Fabian Streit⁷⁸, Patrick F. Sullivan²⁰,²²³,²²⁴, Gustavo Turecki²²⁵, Arne E. Vaaler²²⁶, Eduard Vieta²²⁷, John B. Vincent¹⁰², Irwin D. Waldman²²⁸, Thomas W. Weickert¹⁵²,¹⁷⁶,²¹⁸, Thomas Werge¹⁴,²²⁹,²³⁰,²³¹, Naomi R. Wray¹¹,²³², John-Anker Zwart¹⁸,¹⁹,³³, Joanna M. Biernacka⁸,⁶⁴, John I. Nurnberger²³³, Sven Cichon³,⁴,⁵,⁹⁶, Howard J. Edenberg⁷⁷,²³⁴, Eli A. Stahl¹,²,¹⁸⁰, Andrew McQuillin⁴⁰, Arianna Di Florio⁷⁸,²²⁴, Roel A. Ophoff²³,⁷⁹,¹¹,²³⁵, and Ole A. Andreassen⁶,⁷

See Supplement 1 for all affiliations of consortium members.

## Eating Disorders Working Group of the Psychiatric Genomics Consortium

Roger A.H. Adan¹,²,³, Lars Alfredsson⁴, Tetsuya Ando⁵, Ole A. Andreassen⁶, Harald Aschauer⁷, Jessica M. Baker⁸, Vladimir Bencko⁹, Andrew W. Bergen¹⁰,¹¹, Wade H. Berretti¹², Andreas Birgegård¹³,¹⁴,¹⁵, Joseph M. Boden¹⁶, Ilika Boehm¹⁷, Claudette Boni¹⁸, Vesna Boraska Perica¹⁹,²⁰, Harry Brandt²¹, Gerome Breen²²,²³, Julien Bryois¹⁵, Katharina Buehren²⁴, Cynthia M. Bulik⁸,¹⁵,²⁵, Roland Burghardt²⁶, Laura Carlberg²⁷, Matteo Cassina²⁸, Sven Cichon²⁹,³⁰,³¹, Maurizio Clementi²⁸, Jonathan R.I. Coleman²²,²³, Roger D. Cone³², Philippe Courtet³³, Steven Crawford²¹, Scott Crow³⁴, James J. Crowley¹³,⁵⁹, Unna N. Danner², Oliver S.P. Davis³⁶,³⁷, Martina de Zwaan³⁸, George Dedoussis³⁹, Daniela Degortes⁴⁰, Janiece E. DeSocio⁴¹, Danielle M. Dick⁴²,⁴³,⁴⁴, Dimitris Dikeos⁴⁵, Christian Dina⁶⁰, Monika Dmitrzak-Weglar⁷⁴, Elisa Docampo Martinez⁴⁸,⁴⁹,⁵⁰, Laramie E. Duncan⁵¹, Karin Egberts⁵², Christian R. Marshall¹²⁶, Nicholas G. Martin⁷², Manuel Mattheisen¹³,¹⁴,⁷⁵,¹²⁷, Morten Mattingsdal⁶, Sara McDevitt¹²⁸,¹²⁹, Peter McGuffin²², Sarah E. Medland²⁷, Andres Metspalu⁵³,¹³⁰, Ingrid Meulenbelt¹³¹, Nadia Micali¹³²,¹³³, James Mitchell¹³⁴, Karen Mitchell¹³⁵,¹³⁶, Palmiero Monteleone¹³⁷, Alessio Maria Monteleone¹²⁴, Grant W.
ISGC GWAS of Suicide Attempt

Montgomery<sup>72,86,138</sup>, Preben Bo Mortensen<sup>76,114,115</sup>, Melissa A. Munn-Chernoff<sup>8</sup>, Benedetta Nacmias<sup>139</sup>, Marie Navratilova<sup>63</sup>, Ioanna Ntalla<sup>39</sup>, Catherine M. Olsen<sup>140</sup>, Roel A. Ophoff<sup>141,142</sup>, Julie K. O'Toole<sup>143</sup>, Leonid Padyukov<sup>110</sup>, Aarno Palotie<sup>54,102,144</sup>, Jacques Pantel<sup>18</sup>, Hana Papezova<sup>97</sup>, Richard Parker<sup>72</sup>, John F. Pearson<sup>145</sup>, Nancy L. Pedersen<sup>15</sup>, Stefan Ehrlich<sup>17</sup>, Georgia Escaramis<sup>48,49,50</sup>, Tõnu Esko<sup>53,54</sup>, Thomas Espeseth<sup>55</sup>, Xavier Estivill<sup>48,49,50,56</sup>, Anne Farmer<sup>22</sup>, Angela Favaro<sup>40</sup>, Fernando Fernández-Aranda<sup>57,58</sup>, Manfred M. Fichter<sup>59,60</sup>, Krista Fischer<sup>53</sup>, James A.B. Floyd<sup>61</sup>, Manuel Föcker<sup>62</sup>, Lenka Foretova<sup>63</sup>, Andreas J. Forstner<sup>30,64,65,66</sup>, Monica Forzan<sup>28</sup>, Christopher S. Franklin<sup>19</sup>, Steven Gallinger<sup>67</sup>, Giovanni Gambaro<sup>68</sup>, Héléna A. Gaspar<sup>22,23</sup>, Ina Giegling<sup>69</sup>, Johanna Giuranna<sup>70</sup>, Paola Giusti-Rodriquez<sup>35</sup>, Fragiskos Gonidakis<sup>71</sup>, Scott Gordon<sup>72</sup>, Philip Gorwood<sup>73,74</sup>, Monica Gratacos Mayora<sup>48,49,50</sup>, Jakob Grove<sup>75,76,77,78</sup>, Sébastien Guillaume<sup>33</sup>, Yiran Guo<sup>79</sup>, Hakon Hakonarson<sup>79,80</sup>, Katherine A. Halmi<sup>81</sup>, Ken B. Hanscombe<sup>82</sup>, Konstantinos Hatzikotoulas<sup>19,83</sup>, Joanna Hauser<sup>84</sup>, Johannes Hebebrand<sup>70</sup>, Sietske G. Helder<sup>22,85</sup>, Anjali K. Henders<sup>86</sup>, Stefan Herms<sup>29,30</sup>, Beate Herpertz-Dahlmann<sup>24</sup>, Wolfgang Herzog<sup>87</sup>, Anke Hinney<sup>70</sup>, L. John Horwood<sup>16</sup>, Christopher Hübel<sup>15,22</sup>, Liselotte V. Petersen<sup>76,114,115</sup>, Dilala Pinto<sup>88</sup>, Kirstin L. Purves<sup>22</sup>, Anu Raevuon<sup>101</sup>, Nicolas Ramoz<sup>18</sup>, Ted Reichborn-Kjennerud<sup>112,146</sup>, Valdo Ricca<sup>147</sup>, Samuli Ripatti<sup>148</sup>, Stephan Rippe<sup>149,150,151</sup>, Franziska Ritschel<sup>17,152</sup>, Marion Roberts<sup>22</sup>, Dan Rujescu<sup>69</sup>, Filip Rybakowski<sup>154</sup>, Paolo Santonastaso<sup>155</sup>, André Scherag<sup>156</sup>, Stephen W. Scherer<sup>157,158</sup>, Ulrike Schmidt<sup>22</sup>, Nicholas J. Schork<sup>159</sup>, Alexandra Schosser<sup>160</sup>, Jochen Seitz<sup>24</sup>, Lenka Slachtova<sup>161</sup>, P. Eline Slagboom<sup>131</sup>, Margarita C.T. Slof-Op 't Landt<sup>162,163</sup>, Agnieszka Slopien<sup>164</sup>, Nicole Soranzo<sup>19,165,166,167</sup>, Sandro Sorbi<sup>139,168</sup>, Lorraine Southam<sup>19</sup>, Vidar W. Steen<sup>169,170</sup>, Michael Strober<sup>171,172</sup>, Laura M. Huckins<sup>88</sup>, James I. Hudson<sup>89</sup>, Hartmut Imgart<sup>90</sup>, Hidetoshi Inoko<sup>91</sup>, Vladimir Janout<sup>92</sup>, Susana Jiménez-Murcia<sup>57,58</sup>, Craig Johnson<sup>93</sup>, Jennifer Jordan<sup>94,95</sup>, Antonio Julià<sup>96</sup>, Gursharan Kalsi<sup>22</sup>, Deborah Kaminská<sup>97</sup>, Allan S. Kaplan<sup>98,99,100</sup>, Jaakko Kaprio<sup>101,102</sup>, Leila Karhunen<sup>103</sup>, Andreas Karwautz<sup>104</sup>, Martin J.H. Kas<sup>1,105</sup>, Walter H. Kaye<sup>106</sup>, James L. Kennedy<sup>98,99,100</sup>, Martin A. Kennedy<sup>107</sup>, Anna Kesli-Rahkonen<sup>101</sup>, Kirsty Kiezebrink<sup>108</sup>, Youl-Ri Kim<sup>109</sup>, Katherine M. Kirk<sup>72</sup>, Lars Klareskog<sup>110</sup>, Kelly L. Klump<sup>111</sup>, Gun Peggy S. Knusden<sup>112</sup>, Mikael Landén<sup>15,113</sup>, Janne T. Larsen<sup>76,114,115</sup>, Stephanie Le Hellard<sup>116,117,118</sup>, Virpi M. Leppä<sup>15</sup>, Dong Li<sup>79</sup>, Paul Lichtenstein<sup>15</sup>, Lisa Lilenfeld<sup>119</sup>, Bochao Danae Lin<sup>1</sup>, Jolanta Lissowska<sup>120</sup>, Astri Lundervold<sup>121</sup>, Jurien Luykx<sup>1</sup>, Pierre J. Magistretti<sup>122,123</sup>, Mario Maj<sup>124</sup>, Katrin Mannik<sup>53,125</sup>, Sara Marsal<sup>96</sup>, Garret D. Stuber<sup>8,173</sup>, Patrick F. Sullivan<sup>8,15,35</sup>, Beata Świątkowska<sup>174</sup>, Jin P. Szatkiewicz<sup>35</sup>, Ioanna Tchamazidou<sup>19</sup>, Elena Tenconi<sup>40</sup>, Laura M. Thornton<sup>8</sup>, Alfonso Tortorella<sup>175,176</sup>, Federica Tozzi<sup>177</sup>, Janet Treasure<sup>22</sup>, Artemis Tsitsika<sup>178</sup>, Marta Tyszkiewicz-Nwafor<sup>164</sup>, Konstantinos Tziouvas<sup>179</sup>, Annemarie A. van Elburg<sup>2,180</sup>, Eric F. van Furth<sup>162,163</sup>, Tracey D. Wade<sup>181</sup>, Gudrun Wagner<sup>104</sup>, Esther Walton<sup>17</sup>, Hunna J. Watson<sup>8,182,183</sup>, Thomas Werge<sup>184</sup>, David C. Whiteman<sup>140</sup>, H. Erich Wichmann<sup>185</sup>, Elisabeth Widen<sup>102</sup>, D. Blake Woodside<sup>99,100,186,187</sup>, Shuyang Yao<sup>15</sup>, Zeynep Yilmaz<sup>8,35</sup>, Eleftheria Zeggini<sup>19,83</sup>, Stephanie Zerwas<sup>8</sup>, and Stephan Zipfel<sup>188</sup>

See Supplement 1 for all affiliations of consortium members.

German Borderline Genomics Consortium

Stephanie H. Witt¹, Fabian Streit¹, Martin Jungkunz²,³, Josef Frank¹,
Swapnil Awasthi⁴, Jens Treutlein¹, Lydie Dietl⁵, Cornelia E Schwarze⁶, Nor-
bert Dahmen⁸, Björn H. Schott⁴,⁹, Markus M. Nöthen¹⁰,¹¹, Stephan
Ripke⁴,¹²,¹³, Arian Mobascher⁸, Dan Rujescu⁷, Klaus Lieb⁸, Stefan Roepke⁵,
Christian Schmahl², Martin Bohus³, and Marcella Rietschel¹

See Supplement 1 for all affiliations of consortium members.

MVP Suicide Exemplar Workgroup

Silvia Crivelli, Ph.D. (Lawrence Berkeley National Laboratory), Michelle F. Dennis, B.A. (Durham Veterans Affairs Health Care System & Duke University School of Medicine), Phillip D. Harvey, Ph.D. (University of Miami Miller School of Medicine, Miami, FL), Bruce W. Carter (VA Medical Center), Jennifer E. Huffman, Ph.D. (Massachusetts Veterans Epidemiology Research and Information Center, VA Boston Healthcare System), Daniel Jacobson, Ph.D. (Oak Ridge National Laboratory), Ravi Madduri, Ph.D. (Argonne National Laboratory), John K. Olsen, Ph.D. (Duke University School of Medicine), and John Pestian, Ph.D. (Oak Ridge National Laboratory).

VA Million Veteran Program (MVP)

J. Michael Gaziano, M.D., M.P.H. (co-chair, VA Boston Healthcare System), Sumitra Muralidhar, Ph.D. (co-chair, U.S. Department of

Veterans Affairs), Rachel Ramoni, D.M.D., Sc.D. (U.S. Department of Veterans Affairs), Jean Beckham, Ph.D. (Durham VA Medical Center), Kyong-Mi Chang, M.D. (Philadelphia VA Medical Center), Christopher J. O'Donnell, M.D., M.P.H. (VA Boston Healthcare System), Philip S. Tsao, Ph.D. (VA Palo Alto Health Care System), James Breeling, M.D. (Ex-Officio, U.S. Department of Veterans Affairs), Grant Huang, Ph.D. (Ex-Officio, U.S. Department of Veterans Affairs), and J.H. Casas Romero, M.D., Ph.D. (Ex-Officio, VA Boston Healthcare System). MVP Program Office: Sumitra Muralidhar, Ph.D., and Jennifer Moser, Ph.D., both of U.S. Department of Veterans Affairs. MVP Recruitment/Enrollment: Recruitment/Enrollment Director/Deputy Director, Boston—Stacey B. Whitbourne, Ph.D., Jessica V. Brewer, M.P.H. (VA Boston Healthcare System). MVP Coordinating Centers: Clinical Epidemiology Research Center (CERC), West Haven—Mihaela Aslan, Ph.D. (West Haven VA Medical Center). Cooperative Studies Program Clinical Research Pharmacy Coordinating Center, Albuquerque—Todd Connor, Pharm.D., Dean P. Argyres, B.S., M.S. (New Mexico VA Health Care System). Genetics Coordinating Center, Palo Alto—Philip S. Tsao, Ph.D. (VA Palo Alto Health Care System). MVP Boston Coordinating Center, Boston—J. Michael Gaziano, M.D., M.P.H. (VA Boston Healthcare System). MVP Information Center, Canandaigua—Brady Stephens, M.S. (Canandaigua VA Medical Center). VA Central Biorepository, Boston—Mary T. Brophy, M.D., M.P.H., Donald E. Humphries, Ph.D., Luis E. Selva, Ph.D. (VA Boston Healthcare System). MVP Informatics, Boston—Nhan Do, M.D., Shahpoor Shayan (VA Boston Healthcare System). MVP Data Operations/Analytics, Boston—Kelly Cho, Ph.D. (VA Boston Healthcare System). MVP Science: Science Operations—Christopher J. O'Donnell, M.D., M.P.H. (VA Boston Healthcare System). Genomics Core—Christopher J. O'Donnell, M.D., M.P.H., Saiju Pyarajan, Ph.D. (VA Boston Healthcare System), Philip S. Tsao, Ph.D. (VA Palo Alto Health Care System). Phenomics Core—Kelly Cho, M.P.H., Ph.D. (VA Boston Healthcare System). Data and Computational Sciences—Saiju Pyarajan, Ph.D. (VA Boston Healthcare System). Statistical Genetics—Elizabeth Hauser, Ph.D. (Durham VA Medical Center). Yan Sun, Ph.D. (Atlanta VA Medical Center). Hongyu Zhao, Ph.D. (West Haven VA Medical Center. Current MVP Local Site Investigators: Peter Wilson, M.D. (Atlanta VA Medical Center); Rachel McArdle, Ph.D. (Bay Pines VA Healthcare System); Louis Dellitalia, M.D. (Birmingham VA Medical Center); Kristin Mattocks, Ph.D., M.P.H. (Central Western Massachusetts Healthcare System); John Harley, M.D., Ph.D. (Cincinnati VA Medical Center); Clement J. Zablocki (VA Medical Center); Jeffrey Whittle, M.D., M.P.H.; Frank Jacono, M.D. (VA Northeast Ohio Healthcare System); Jean Frank, Ph.D. (Durham VA Medical Center); Edith Nourse Rogers Memorial Veterans Hospital; Salvador Gutierrez, M.D. (Edward Hines, Jr. VA Medical Center); Gretchen Gibson, D.D.S., M.P.H. (Veterans Health Care System of the Ozarks); Kimberly Hammer, Ph.D. (Fargo VA Health Care System); Laurence Kaminsky, Ph.D. (VA Health Care Upstate New York); Gerardo Villareal, M.D. (New Mexico VA Health Care System); Scott Kinlay, M.B.B.S., Ph.D. (VA Boston Healthcare System); Junzhe Xu, M.D. (VA Western New York Healthcare System); Mark Hamner, M.D. (Ralph H. Johnson VA Medical Center); Roy Mathew, M.D. (Columbia VA Health Care System); Sujata Bhushan, M.D. (VA North Texas Health Care System); Pran Iuvanti, DO, Ph.D. (Hampton VA Medical Center); Michael Godschalk, M.D. (Richmond VA Medical Center); Zuhair Ballas, M.D. (Iowa City VA Health Care System); Douglas Ivins, M.D. (Eastern Oklahoma VA Health Care System); Stephen Mastorides, M.D. (James A. Haley Veterans' Hospital); Jonathan Moorman, M.D., Ph.D. (James H. Quillen VA Medical Center); Saib Gappy, M.D. (John D. Dingell VA Medical Center); Jon Klein, M.D., Ph.D. (Louisville VA Medical Center); Nora Ratcliffe, M.D. (Manchester VA Medical Center); Hermes Florez, M.D., Ph.D. (Miami VA Health Care System); Olaoluwa Okusaga, M.D. (Michael E. DeBakey VA Medical Center); Maureen Murdoch, M.D., M.P.H. (Minneapolis VA Health Care System); Peruvema Sriram, M.D. (N FL/S GA Veterans Health System); Shing Shing Yeh, Ph.D., M.D. (Northport VA Medical Center); Neeraj Tandon, M.D. (Overton Brooks VA Medical Center); Darshana Jhala, M.D. (Philadelphia VA Medical Center); Samuel Aguayo, M.D. (Phoenix VA Health Care System); David Cohen, M.D. (Portland VA Medical Center); Satish Sharma, M.D. (Providence VA Medical Center); Suthat Liangpunskul, M.D., M.P.H. (Richard Roudebush VA Medical Center); Kris Ann Oursler, M.D. (Salem VA
Medical Center); Mary Whooley, M.D. (San Francisco VA Health Care System); Sunil Ahuja, M.D. (South Texas Veterans Health Care System); Joseph Constans, Ph.D. (Southeast Louisiana Veterans Health Care System); Paul Meyer, M.D., Ph.D. (Southern Arizona VA Health Care System); Jennifer Greco, M.D. (Sioux Falls VA Health Care System); Michael Rauchman, M.D. (St. Louis VA Health Care System); Richard Servatius, Ph.D. (Syracuse VA Medical Center); Melinda Gaddy, Ph.D. (VA Eastern Kansas Health Care System); Agnes Wallbom, M.D., M.S. (VA Greater Los Angeles Health Care System); Timothy Morgan, M.D. (VA Long Beach Healthcare System); Todd Stapley, D.O. (VA Maine Healthcare System); Scott Sherman, M.D., M.P.H. (VA New York Harbor Healthcare System); George Ross, M.D. (VA Pacific Islands Health Care System); Philip Tsao, Ph.D. (VA Palo Alto Health Care System); Patrick Strollo Jr., M.D. (VA Pittsburgh Health Care System); Edward Boyko, M.D. (VA Puget Sound Health Care System); Laurence Meyer, M.D., Ph.D. (VA Salt Lake City Health Care System); Samir Gupta, M.D., M.S.C.S. (VA San Diego Healthcare System); Mostaqul Huq, Pharm.D., Ph.D. (VA Sierra Nevada Health Care System); Joseph Fayad, M.D. (VA Southern Nevada Healthcare System); Adriana Hung, M.D., M.P.H. (VA Tennessee Valley Healthcare System); Jack Lichy, M.D., Ph.D. (Washington, DC VA Medical Center); Robin Hurley, M.D. (W.G., Bill Hefner VA Medical Center); Brooks Robey, M.D. (White River Junction VA Medical Center); and Robert Striker, M.D., Ph.D. (William S. Middleton Memorial Veterans Hospital).

NM and JK contributed equally to this work.

Address correspondence to Niamh Mullins, Ph.D., at niamh.mullins@mssm.edu, or Douglas M. Ruderfer, Ph.D., at douglas.ruderfer@vanderbilt.edu.

Received Jan 5, 2021; revised May 7, 2021; accepted May 26, 2021.

Supplementary material cited in this article is available online at https://doi.org/10.1016/j.biopsych.2021.05.029.

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