About the Authors

Lily Paemka

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Program in Genetics, The University of Iowa, Iowa City, Iowa, United States of America

Vinit B. Mahajan

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Ophthalmology and Visual Sciences, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America

Jessica M. Skeie

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Ophthalmology and Visual Sciences, The University of Iowa, Iowa City, Iowa, United States of America

Levi P. Sowers

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Neurology, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America

Salleh N. Ehaideb

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Program in Genetics, The University of Iowa, Iowa City, Iowa, United States of America, Department of Biology, The University of Iowa, Iowa City, Iowa, United States of America

Pedro Gonzalez-Alegre

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Neurology, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Graduate Program of Neuroscience, The University of Iowa, Iowa City, Iowa, United States of America

Toshikuni Sasaoka

Affiliation Center for Bioresources, Brain Research Institute, Niigata University, Niigata, Japan

Hirotaka Tao

Affiliation Hospital for Sick Kids, University of Toronto, Toronto, Canada

Asuka Miyagi

Affiliation Developmental Biology Department, National Institute for Basic Biology, Okazaki City, Japan

Naoto Ueno

Affiliation Developmental Biology Department, National Institute for Basic Biology, Okazaki City, Japan

Keizo Takao

Affiliations Section of Behavior Patterns, Center for Genetic Analysis of Behavior National Institute for Physiological Sciences, Okazaki, Japan, Japan Science and Technology Agency, Kawaguchi-shi, Japan

Tsuyoshi Miyakawa

Affiliations Section of Behavior Patterns, Center for Genetic Analysis of Behavior National Institute for Physiological Sciences, Okazaki, Japan, Japan Science and Technology Agency, Kawaguchi-shi, Japan, Division of Systems Medical Science, Institute for Comprehensive Medical Science, Fujita Health University, Toyoake, Japan

Shu Wu

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America

Benjamin W. Darbro

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Department of Pathology, The University of Iowa, Iowa City, Iowa, United States of America

Polly J. Ferguson

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America

Andrew A. Pieper

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Neurology, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Department of Psychiatry, The University of Iowa, Iowa City, Iowa, United States of America

Jeremiah K. Britt

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Department of Psychiatry, The University of Iowa, Iowa City, Iowa, United States of America

John A. Wemmie

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Graduate Program of Neuroscience, The University of Iowa, Iowa City, Iowa, United States of America, Department of Psychiatry, The University of Iowa, Iowa City, Iowa, United States of America

Danielle S. Rudd

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Program in Genetics, The University of Iowa, Iowa City, Iowa, United States of America, Department of Psychiatry, The University of Iowa, Iowa City, Iowa, United States of America

Thomas Wassink

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Department of Psychiatry, The University of Iowa, Iowa City, Iowa, United States of America

Hatem El-Shanti

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Shafallah Medical Genetics Center, Doha, Qatar

Heather C. Mefford

Affiliation Division of Genetic Medicine, Department of Pediatrics, University of Washington, Seattle, Washington, United States of America

Gemma L. Carvill

Affiliation Division of Genetic Medicine, Department of Pediatrics, University of Washington, Seattle, Washington, United States of America

J. Robert Manak

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Program in Genetics, The University of Iowa, Iowa City, Iowa, United States of America, Department of Biology, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Graduate Program in Molecular and Cellular Biology, The University of Iowa, Iowa City, Iowa, United States of America

Alexander G. Bassuk

alexander-bassuk@uiowa.edu

Affiliations The University of Iowa, Iowa City, Iowa, United States of America, Department of Pediatrics, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Program in Genetics, The University of Iowa, Iowa City, Iowa, United States of America, Department of Neurology, The University of Iowa, Iowa City, Iowa, United States of America, Roy J. and Lucille A. Carver College of Medicine, The University of Iowa, Iowa City, Iowa, United States of America, Interdisciplinary Graduate Program in Molecular and Cellular Biology, The University of Iowa, Iowa City, Iowa, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: LP PG NU KT JRM AGB TM. Performed the experiments: LP SW JMS SNE HT AM KT TM LPS. Analyzed the data: LP TW DSR AAP JMS HCM GLC LPS KT JAW PJF HE BWD TS HT JKB TM. Contributed reagents/materials/analysis tools: AGB VBM PG KT TM JRM. Wrote the paper: LP VBM JRM AGB.