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Published Online: 1 September 2011

Analysis of 94 Candidate Genes and 12 Endophenotypes for Schizophrenia From the Consortium on the Genetics of Schizophrenia

Tiffany A. Greenwood, Ph.D., Laura C. Lazzeroni, Ph.D., Sarah S. Murray, Ph.D., Kristin S. Cadenhead, M.D., Monica E. Calkins, Ph.D., Dorcas J. Dobie, M.D., Michael F. Green, Ph.D., Raquel E. Gur, M.D., Ph.D., Ruben C. Gur, Ph.D., Gary Hardiman, Ph.D., John R. Kelsoe, M.D., Sherry Leonard, Ph.D., Gregory A. Light, Ph.D., Keith H. Nuechterlein, Ph.D., Ann Olincy, M.D., Allen D. Radant, M.D., Nicholas J. Schork, Ph.D., Larry J. Seidman, Ph.D., Larry J. Siever, M.D., Jeremy M. Silverman, Ph.D., William S. Stone, Ph.D., Neal R. Swerdlow, M.D., Ph.D., Debby W. Tsuang, M.D., M.Sc., Ming T. Tsuang, M.D., Ph.D., D.Sc., Bruce I. Turetsky, M.D., Robert Freedman, M.D., and David L. Braff, M.D.Authors Info & Affiliations
Publication: American Journal of Psychiatry
Volume 168, Number 9
https://doi.org/10.1176/appi.ajp.2011.10050723
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Abstract

Genes affecting glutamate neurotransmission featured prominently in associations between 94 genes and 12 inherited physiological or cognitive characteristics of schizophrenia. Of 16,620 possible SNP-trait associations, 47 showed strong significance.

Abstract

Objective:

The authors used a custom array of 1,536 single-nucleotide polymorphisms (SNPs) to interrogate 94 functionally relevant candidate genes for schizophrenia and identify associations with 12 heritable neurophysiological and neurocognitive endophenotypes in data collected by the Consortium on the Genetics of Schizophrenia.

Method:

Variance-component association analyses of 534 genotyped subjects from 130 families were conducted by using Merlin software. A novel bootstrap total significance test was also developed to overcome the limitations of existing genomic multiple testing methods and robustly demonstrate significant associations in the context of complex family data and possible population stratification effects.

Results:

Associations with endophenotypes were observed for 46 genes of potential functional significance, with three SNPs at p<10–4, 27 SNPs at p<10–3, and 147 SNPs at p<0.01. The bootstrap analyses confirmed that the 47 SNP-endophenotype combinations with the strongest evidence of association significantly exceeded that expected by chance alone, with 93% of these findings expected to be true. Many of the genes interact on a molecular level, and eight genes (e.g., NRG1 and ERBB4) displayed evidence for pleiotropy, revealing associations with four or more endophenotypes. The results collectively support a strong role for genes related to glutamate signaling in mediating schizophrenia susceptibility.

Conclusions:

This study supports use of relevant endophenotypes and the bootstrap total significance test for identifying genetic variation underlying the etiology of schizophrenia. In addition, the observation of extensive pleiotropy for some genes and singular associations for others suggests alternative, independent pathways mediating pathogenesis in the “group of schizophrenias.”

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Supplementary Material

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Information & Authors

Information

Published In

Go to American Journal of Psychiatry
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American Journal of Psychiatry
Volume 168 • Number 9 • September 2011
Pages: 930 - 946
PubMed: 21498463

History

Received: 17 May 2010
Revision received: 12 October 2010
Revision received: 20 December 2010
Accepted: 4 February 2011
Published online: 1 September 2011
Published in print: September 2011

Authors

Affiliations

Tiffany A. Greenwood, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Laura C. Lazzeroni, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Sarah S. Murray, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Kristin S. Cadenhead, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Monica E. Calkins, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Dorcas J. Dobie, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Michael F. Green, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Raquel E. Gur, M.D., Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Ruben C. Gur, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Gary Hardiman, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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John R. Kelsoe, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Sherry Leonard, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Gregory A. Light, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Keith H. Nuechterlein, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Ann Olincy, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Allen D. Radant, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Nicholas J. Schork, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Larry J. Seidman, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Larry J. Siever, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Jeremy M. Silverman, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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William S. Stone, Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Neal R. Swerdlow, M.D., Ph.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Debby W. Tsuang, M.D., M.Sc.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Ming T. Tsuang, M.D., Ph.D., D.Sc.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Bruce I. Turetsky, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Robert Freedman, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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David L. Braff, M.D.
From the Department of Psychiatry, the Department of Medicine, and the Biomedical Genomics Laboratory, University of California at San Diego, La Jolla; the San Diego VA Healthcare System; Scripps Genomic Medicine and the Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, Calif.; the Department of Psychiatry and Behavioral Sciences and the Department of Pediatrics, Stanford University, Stanford, Calif.; the Department of Psychiatry, University of Pennsylvania, Philadelphia; the Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle; the VA Puget Sound Health Care System, Seattle; the Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, UCLA; the Department of Psychiatry, University of Colorado Health Sciences Center, Denver; the Department of Psychiatry, Harvard Medical School and the Harvard Institute of Psychiatric Epidemiology and Genetics, Boston; the Mental Health Center Public Psychiatry Division, Beth Israel Deaconess Medical Center, Boston; the Department of Psychiatry, Mount Sinai School of Medicine, New York; and the James J. Peters VA Medical Center, New York.
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Notes

Address correspondence and reprint requests to Dr. Braff, Department of Psychiatry, University of California at San Diego, Mail Code 0804, 9500 Gilman Dr., La Jolla, CA 92093-0804; [email protected] (e-mail).

Funding Information

Dr. Greenwood has received unrelated support for consulting services from INFOTECH Soft. Dr. Freedman has a patent through the VA on DNA sequences in CHRNA7. Dr. Green has been a consultant for Abbott, Cypress, Dainippon Sumitomo, Lundbeck, Otsuka, Sanofi-Aventis, Takeda, and Teva, and he has been a speaker for Janssen Cilag, Otsuka, and Sunovion. Dr. Raquel Gur has received unrelated research support from AstraZeneca and Pfizer. Dr. Ruben Gur is a consultant to Johnson and Johnson and has received investigator-initiated grants from AstraZeneca, Merck, and Pfizer. Dr. Kelsoe is a consultant for AstraZeneca and Psynomics and is on a speakers bureau for Merck. Dr. Light has received unrelated support for consulting services from Allergan, Memory, and Roche. Dr. Nuechterlein reports an investigator-initiated research grant from Ortho-McNeil Janssen Scientific Affairs and consultation to Merck and Wyeth. Dr. Olincy has received unrelated research support from Lundbeck. Dr. Turetsky has received unrelated research support from AstraZeneca and Pfizer and is a consultant to Roche. The other authors report no financial relationships with commercial interests.This study was supported by NIMH grants R01-MH-065571, R01-MH-065588, R01-MH-065562, R01-MH-065707, R01-MH-065554, R01-MH-065578, R01-MH-065558, R01-MH-86135, P50-MH-086383 (Dr. Freedman), and K01-MH-087889 (Dr. Greenwood); by grant DK-063491 from the National Institute of Diabetes and Digestive and Kidney Diseases (Dr. Hardiman); by the Department of Veterans Affairs, Veterans Integrated Service Network (VISN) 22, Mental Illness Research, Education, and Clinical Center (Dr. Braff); by a NARSAD Distinguished Investigator Award (Dr. Braff); and by additional salary support of Dr. Lazzeroni from VA Cooperative Studies Program study 478 and the Department of Veterans Affairs VISN-21 Mental Illness Research, Education, and Clinical Center.

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