Cross-cohort analysis of expression and splicing quantitative trait loci in TOPMed

P Peter Orchard (Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA.) T Thomas W. Blackwell (Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.) L Linda Kachuri (Department of Epidemiology and Population Health, School of Medicine, Stanford University, Stanford, CA, USA.) P Peter J. Castaldi (Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.) M Michael H. Cho (Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.) S Stephanie A. Christenson (Division of Pulmonary, Critical Care, Allergy, and Sleep Medicine, School of Medicine, University of California, San Francisco, San Francisco, CA, USA.) P Peter Durda (Department of Pathology and Laboratory Medicine, Larner College of Medicine, University of Vermont, Burlington, VT, USA.) S Stacey Gabriel C Craig P. Hersh (Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.) S Scott Huntsman (Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.) S Seungyong Hwang (Department of Genetics, Stanford University, Stanford, CA, USA.) R Roby Joehanes (Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.) M Mari Johnson (Division of Public Health Sciences, Fred Hutchinson Cancer Center, Seattle, WA, USA.) X Xingnan Li (Division of Data Driven and Digital Medicine, Department of Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA.) H Honghuang Lin (Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.) C Ching-Ti Liu (Department of Biostatistics, School of Public Health, Boston University, Boston, MA, USA.) Y Yongmei Liu (Division of Cardiology, Department of Medicine, School of Medicine, Duke University, Durham, NC, USA.) A Angel C. Y. Mak (Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.) A Ani W. Manichaikul (Department of Genome Sciences, School of Medicine, University of Virginia, Charlottesville, VA, USA.) D David T. Paik A Aabida Saferali (Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.) J Joshua D. Smith K Kent D. Taylor (The Institute for Translational Genomics and Population Sciences, Department of Pediatrics, The Lundquist Institute for Biomedical Innovation at Harbor-UCLA Medical Center, Torrance, CA, USA.) R Russell P. Tracy (Laboratory for Clinical Biochemistry Research, Larner College of Medicine, University of Vermont, Burlington, VT, USA.) J Jiongming Wang (Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.) M Mingqiang Wang J Joshua S. Weinstock (Department of Biomedical Engineering, Johns Hopkins University, Baltimore, MD, USA.) J Jeffrey Weiss H Heather E. Wheeler (Program in Bioinformatics, Loyola University Chicago, Chicago, IL, USA.) Y Ying Zhou S Sebastian Zöllner (Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.) J Joseph C. Wu L Luisa Mestroni (Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.) S Sharon Graw (Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.) M Matthew R. G. Taylor (Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.) V Victor E. Ortega (Department of Medicine, Division of Pulmonary and Critical Care Medicine, Mayo Clinic, Phoenix, AZ, USA.) W W. Craig Johnson (Collaborative Health Studies Coordinating Center, University of Washington, Seattle, WA, USA.) W Weiniu Gan (National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.) G Gonçalo Abecasis D Deborah A. Nickerson (Northwest Genomics Center, University of Washington, Seattle, WA, USA.) N Namrata Gupta (Broad Institute of MIT and Harvard, Cambridge, MA, USA.) K Kristin Ardlie (Broad Institute of MIT and Harvard, Cambridge, MA, USA.) P Prescott G. Woodruff (Division of Pulmonary, Critical Care, Allergy, and Sleep Medicine, School of Medicine, University of California, San Francisco, San Francisco, CA, USA.) R Russell P. Bowler (Department of Genomic Sciences and Systems Biology, Cleveland Clinic, Cleveland, OH, USA.) D Deborah A. Meyers (Department of Medicine, Mayo Clinic, Scottsdale, AZ, USA.) A Alex Reiner (Department of Epidemiology, University of Washington, Seattle, WA, USA.) C Charles Kooperberg (Division of Public Health Sciences, Fred Hutchinson Cancer Center, Seattle, WA, USA.) E Elad Ziv (Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.) R Ramachandran S. Vasan (Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.) M Martin G. Larson (Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.) L L. Adrienne Cupples (Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.) E Edwin K. Silverman (Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.) S Stephen S. Rich (Department of Genome Sciences, School of Medicine, University of Virginia, Charlottesville, VA, USA.) N Nancy Heard-Costa (Chobanian & Avedisian School of Medicine, Boston University, Boston, MA, USA.) H Hua Tang (Department of Genetics, Stanford University, Stanford, CA, USA.) J Jerome I. Rotter A Albert V. Smith (Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.) D Daniel Levy (Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.) F François Aguet L Laura J. Scott L Laura M. Raffield (Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.) S Stephen C. J. Parker (Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA.) N Namiko Abe (New York Genome Center, New York, NY, USA.) L Laura Almasy (Children’s Hospital of Philadelphia, University of Pennsylvania, Philadelphia, PA, USA.) S Seth Ament (University of Maryland, Baltimore, MD, USA.) P Pramod Anugu (University of Mississippi, Jackson, MS, USA.) P Paul Auer (Medical College of Wisconsin, Milwaukee, WI, USA.) D Dimitrios Avramopoulos (Johns Hopkins University, Baltimore, MD, USA.) A Adithya Balasubramanian (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) R R. Graham Barr (Columbia University, New York, NY, USA.) L Lucas Barwick (The Emmes Corporation, Rockville, MD, USA.) T Terri Beaty (Johns Hopkins University, Baltimore, MD, USA.) D Diane Becker (Johns Hopkins University, Baltimore, MD, USA.) L Lewis Becker (Johns Hopkins University, Baltimore, MD, USA.) A Amber Beitelshees (University of Maryland, Baltimore, MD, USA.) T Takis Benos (University of Pittsburgh, Pittsburgh, PA, USA.) M Marcos Bezerra (Fundação de Hematologia e Hemoterapia de Pernambuco - Hemope, Recife, Brazil.) J Joshua Bis (University of Washington, Seattle, WA, USA.) J Jennifer Brody (University of Washington, Seattle, WA, USA.) U Ulrich Broeckel (Medical College of Wisconsin, Milwaukee, WI, USA.) J Jai Broome (University of Washington, Seattle, WA, USA.) K Karen Bunting (New York Genome Center, New York, NY, USA.) E Erin Buth (University of Washington, Seattle, WA, USA.) V Vincent Carey (Brigham andWomen’s Hospital, Boston, MA, USA.) C Cara Carty (Washington State University, Pullman, WA, USA.) R Richard Casaburi (University of California, Los Angeles, Los Angeles, CA, USA.) M Mark Chaffin (Broad Institute, Cambridge, MA, USA.) C Christy Chang (University of Maryland, Baltimore, MD, USA.) Y Yi-Cheng Chang (National Taiwan University, Taipei, Taiwan (Province of China).) S Sameer Chavan (University of Colorado at Denver, Denver, CO, USA.) B Bo-Juen Chen (New York Genome Center, New York, NY, USA.) W Wei-Min Chen (University of Virginia, Charlottesville, VA, USA.) S Seung Hoan Choi (Broad Institute, Cambridge, MA, USA.) L Lee-Ming Chuang (National Taiwan University, Taipei, Taiwan (Province of China).) R Ren-Hua Chung (National Health Research Institute Taiwan, Miaoli County, Taiwan (Province of China).) M Matthew Conomos (University of Washington, Seattle, WA, USA.) E Elaine Cornell (University of Vermont, Burlington, VT, USA.) C Carolyn Crandall (University of California, Los Angeles, Los Angeles, CA, USA.) J James Crapo (National Jewish Health, Denver, CO, USA.) J Jeffrey Curtis (University of Michigan, Ann Arbor, MI, USA.) C Coleen Damcott (University of Maryland, Baltimore, MD, USA.) S Sean David (University of Chicago, Chicago, IL, USA.) L Lisa de las Fuentes (Washington University in St. Louis, St. Louis, MO, USA.) P Paul de Vries (University of Texas Health at Houston, Houston, TX, USA.) R Ranjan Deka (University of Cincinnati, Cincinnati, OH, USA.) D Dawn DeMeo (Brigham andWomen’s Hospital, Boston, MA, USA.) S Scott Devine (University of Maryland, Baltimore, MD, USA.) H Huyen Dinh (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) H Harsha Doddapaneni Q Qing Duan (University of North Carolina, Chapel Hill, NC, USA.) R Ravi Duggirala (University of Texas Rio Grande Valley School of Medicine, Edinburg, TX, USA.) C Charles Eaton (Harvard University, Cambridge, MA, USA.) L Lynette Ekunwe (University of Mississippi, Jackson, MS, USA.) A Adel El Boueiz (Washington University in St. Louis, St. Louis, MO, USA.) L Leslie Emery (University of Washington, Seattle, WA, USA.) C Charles Farber (University of Virginia, Charlottesville, VA, USA.) J Jesse Farek (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) N Nora Franceschini (University of North Carolina, Chapel Hill, NC, USA.) C Chris Frazar (University of Washington, Seattle, WA, USA.) M Mao Fu (University of Maryland, Baltimore, MD, USA.) S Stephanie M. Fullerton (University of Washington, Seattle, WA, USA.) L Lucinda Fulton (Washington University in St. Louis, St. Louis, MO, USA.) S Shanshan Gao (University of Colorado at Denver, Denver, CO, USA.) Y Yan Gao M Margery Gass (Fred Hutchinson Cancer Research Center, Seattle, WA, USA.) H Heather Geiger A Auyon Ghosh (Brigham andWomen’s Hospital, Boston, MA, USA.) C Chris Gignoux (Stanford University, Stanford, CA, USA.) D David Glahn (Boston Children’s Hospital, Harvard Medical School, Boston, MA, USA.) S Stephanie Gogarten (University of Washington, Seattle, WA, USA.) D Da-Wei Gong (University of Maryland, Baltimore, MD, USA.) H Harald Goring (University of Texas Rio Grande Valley School of Medicine, San Antonio, TX, USA.) D Daniel Grine (University of Colorado at Denver, Denver, CO, USA.) C C. Charles Gu (Washington University in St. Louis, St. Louis, MO, USA.) Y Yue Guan (University of Maryland, Baltimore, MD, USA.) M Michael Hall (University of Mississippi, Jackson, MS, USA.) Y Yi Han D Daniel Harris (University of Maryland, Philadelphia, PA, USA.) B Ben Heavner D David Herrington (Wake Forest Baptist Health, Winston-Salem, NC, USA.) B Brian Hobbs E Elliott Hong (University of Maryland, Baltimore, MD, USA.) K Karin Hoth (University of Iowa, Iowa City, IA.) C Chao (Agnes) Hsiung (National Health Research Institute Taiwan, Miaoli County, Taiwan (Province of China).) J Jianhong Hu Y Yi-Jen Hung (Tri-Service General Hospital National Defense Medical Center, Taiwan (Province of China).) H Haley Huston (Blood Works Northwest, Seattle, WA, USA.) C Chii Min Hwu (Taichung Veterans General Hospital Taiwan, Taichung City, Taiwan (Province of China).) R Rebecca Jackson D Deepti Jain (University of Washington, Seattle, WA, USA.) J Jill Johnsen (University of Washington, Seattle, WA, USA.) R Rich Johnston (Emory University, Atlanta, GA, USA.) K Kimberly Jones (Johns Hopkins University, Baltimore, MD, USA.) M Michael Kessler A Alyna Khan (University of Washington, Seattle, WA, USA.) Z Ziad Khan W Wonji Kim (Harvard University, Cambridge, MA, USA.) J John Kimoff (McGill University, Montréal, QC, Canada.) G Greg Kinney (University of Colorado at Denver, Aurora, CO, USA.) H Holly Kramer (Loyola University, Maywood, IL, USA.) C Christoph Lange (Harvard School of Public Health, Boston, MA, USA.) E Ethan Lange (University of Colorado at Denver, Denver, CO, USA.) C Cathy Laurie (University of Washington, Seattle, WA, USA.) C Cecelia Laurie (University of Washington, Seattle, WA, USA.) M Meryl LeBoff (Brigham andWomen’s Hospital, Boston, MA, USA.) S Sandra Lee (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) W Wen-Jane Lee (Taichung Veterans General Hospital Taiwan, Taichung City, Taiwan (Province of China).) D David Levine (University of Washington, Seattle, WA, USA.) J Joshua Lewis (University of Maryland, Baltimore, MD, USA.) Y Yun Li X Xihong Lin S Simin Liu (Brown University, Providence, RI, USA.) Y Yu Liu B Barry Make (Johns Hopkins University, Baltimore, MD, USA.) A Alisa Manning (Broad Institute, Harvard University, Massachusetts General Hospital, Boston, MA, USA.) J JoAnn Manson (Brigham andWomen’s Hospital, Boston, MA, USA.) L Lisa Martin M Melissa Marton (New York Genome Center, New York City, NY, USA.) S Susan Mathai (University of Colorado at Denver, Denver, CO, USA.) S Susanne May P Patrick McArdle (University of Maryland, Baltimore, MD, USA.) M Merry-Lynn McDonald S Sean McFarland (Harvard University, Cambridge, MA, USA.) D Daniel McGoldrick (University of Washington, Seattle, WA, USA.) C Caitlin McHugh (University of Washington, Seattle, WA, USA.) H Hao Mei (University of Mississippi, Jackson, MS, USA.) J James Meigs (Massachusetts General Hospital, Boston, MA, USA.) V Vipin Menon (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) N Nancy Min (University of Mississippi, Jackson, MS, USA.) M Matt Moll (Brigham andWomen’s Hospital, Boston, MA, USA.) Z Zeineen Momin (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) M May Montasser (National Heart, Lung, and Blood Institute, Bethesda, MD, USA.) J Josyf C Mychaleckyj (University of Virginia, Charlottesville, VA, USA.) R Rakhi Naik (Johns Hopkins University, Baltimore, MD, USA.) T Take Naseri P Pradeep Natarajan S Sarah C. Nelson B Bonnie Neltner (University of Colorado at Denver, Denver, CO, USA.) C Caitlin Nessner (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) O Osuji Nkechinyere (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) J Jeff O'Connell (University of Maryland, Baltimore, MD, USA.) T Tim O’Connor (University of Maryland, Baltimore, MD, USA.) H Heather Ochs-Balcom (University at Buffalo, Buffalo, NY, USA.) G Geoffrey Okwuonu (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) J James Pankow (University of Minnesota, Minneapolis, MN, USA.) C Cora Parker (RTI International, Research Triangle Park, NC, USA.) G Gina Peloso (Boston University, Boston, MA, USA.) J Juan Manuel Peralta (University of Texas Rio Grande Valley School of Medicine, Edinburg, TX, USA.) M Marco Perez (Stanford University, Stanford, CA, USA.) J James Perry (University of Maryland, Baltimore, MD, USA.) U Ulrike Peters (Fred Hutchinson Cancer Research Center, Seattle, WA, USA.) L Lawrence S Phillips (Emory University, Atlanta, GA, USA.) T Toni Pollin (University of Maryland, Baltimore, MD, USA.) J Julia Powers Becker (University of Colorado at Denver, Denver, CO, USA.) M Meher Preethi Boorgula (University of Colorado at Denver, Denver, CO, USA.) B Bruce Psaty (University of Washington, Seattle, WA, USA.) D Dandi Qiao (Brigham andWomen’s Hospital, Boston, MA, USA.) N Nicholas Rafaels (University of Colorado at Denver, Denver, CO, USA.) M Mahitha Rajendran (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) L Laura Rasmussen-Torvik (Northwestern University, Chicago, IL, USA.) A Aakrosh Ratan (University of Virginia, Charlottesville, VA, USA.) R Robert Reed (University of Maryland, Baltimore, MD, USA.) E Elizabeth Regan (National Jewish Health, Denver, CO, USA.) M Muagututi‘a Sefuiva Reupena (Lutia I Puava Ae Mapu I Fagalele, Apia, Samoa.) R Rebecca Robillard (University of Ottawa, Ottawa, ON, Canada.) C Carolina Roselli (Broad Institute, Cambridge, MA, USA.) I Ingo Ruczinski (Johns Hopkins University, Baltimore, MD, USA.) A Alexi Runnels P Pamela Russell (University of Colorado at Denver, Denver, CO, USA.) K Kathleen Ryan (University of Maryland, Baltimore, MD, USA.) E Ester Cerdeira Sabino (Universidade de Sao Paulo, Sao Paulo, Brazil.) S Shabnam Salimi (University of Maryland, Seattle, WA, USA.) S Sejal Salvi (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) S Steven Salzberg (Johns Hopkins University, Baltimore, MD, USA.) K Kevin Sandow (Lundquist Institute, Torrance, CA, USA.) J Jireh Santibanez (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) K Karen Schwander (Washington University in St. Louis, St. Louis, MO, USA.) F Frank Sciurba (University of Pittsburgh, Pittsburgh, PA, USA.) F Frédéric Sériès (Université Laval, Quebec City, Canada.) A Amol Shetty (University of Maryland, Baltimore, MD, USA.) A Aniket Shetty (University of Colorado at Denver, Denver, CO, USA.) B Brian Silver (UMass Memorial Medical Center, Worcester, MA, USA.) R Robert Skomro (University of Saskatchewan, Saskatoon, Canada.) T Tanja Smith (New York Genome Center, New York, NY, USA.) S Sylvia Smoller (Albert Einstein College of Medicine, New York, NY, USA.) B Beverly Snively (Wake Forest Baptist Health, Winston-Salem, NC, USA.) A Adrienne M. Stilp G Garrett Storm (University of Colorado at Denver, Aurora, CO, USA.) E Elizabeth Streeten (University of Maryland, Baltimore, MD, USA.) J Jessica Lasky Su (Brigham andWomen’s Hospital, Boston, MA, USA.) Y Yun Ju Sung (Washington University in St. Louis, St. Louis, MO, USA.) J Jody Sylvia (Brigham andWomen’s Hospital, Boston, MA, USA.) A Adam Szpiro (University of Washington, Seattle, WA, USA.) M Margaret Taub (Johns Hopkins University, Baltimore, MD, USA.) S Simeon Taylor (University of Maryland, Baltimore, MD, USA.) T Timothy A. Thornton (University of Washington, Seattle, WA, USA.) M Machiko Threlkeld (University of Washington, Seattle, WA, USA.) L Lesley Tinker (Fred Hutchinson Cancer Research Center, Seattle, WA, USA.) D David Tirschwell (University of Washington, Seattle, WA, USA.) H Hemant Tiwari (University of Alabama, Birmingham, AL, USA.) C Catherine Tong (University of Washington, Seattle, WA, USA.) M Michael Tsai (University of Minnesota, Minneapolis, MN, USA.) D Dhananjay Vaidya (Johns Hopkins University, Baltimore, MD, USA.) T Tarik Walker (University of Colorado at Denver, Denver, CO, USA.) R Robert Wallace (University of Iowa, Iowa City, IA.) A Avram Walts (University of Colorado at Denver, Denver, CO, USA.) F Fei Fei Wang (University of Washington, Seattle, WA, USA.) H Heming Wang K Karol Watson (University of California, Los Angeles, Los Angeles, CA, USA.) J Jennifer Watt (Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.) L Lu-Chen Weng (Massachusetts General Hospital, Boston, MA, USA.) J Jennifer Wessel (Indiana University, Indianapolis, IN, USA.) K Kayleen Williams (University of Washington, Seattle, WA, USA.) C Carla Wilson (Brigham andWomen’s Hospital, Boston, MA, USA.) J James Wilson L Lara Winterkorn (New York Genome Center, New York City, NY, USA.) Q Quenna Wong B Baojun Wu (Henry Ford Health System, Detroit, MI, USA.) H Huichun Xu (University of Maryland, Baltimore, MD, USA.) L Lisa Yanek (Johns Hopkins University, Baltimore, MD, USA.) I Ivana Yang (University of Colorado at Denver, Denver, CO, USA.) S Seyedeh Maryam Zekavat (Broad Institute, Cambridge, MA, USA.) S Snow Xueyan Zhao (National Jewish Health, Denver, CO, USA.) W Wei Zhao X Xiaofeng Zhu

Abstract

Most genetic variants associated with complex traits are hypothesized to regulate gene expression. To understand the genetics underlying gene expression variability, we characterized 14,324 RNA-sequencing samples from the Trans-Omics for Precision Medicine program and performed expression and splicing quantitative trait locus (e/sQTL) analyses in six tissues and cell types, including whole blood ( n = 6454) and lung ( n = 1291). We detected tens of thousands of secondary cis-e/sQTLs, showing that secondary cis-e/sQTL discovery remains unsaturated. We fine-mapped UK Biobank–derived genome-wide association study (GWAS) signals from 164 traits and identified e/sQTL colocalizations for 10,611 GWAS signals, including 7096 that colocalize with secondary e/sQTLs. Our results suggest that even larger e/sQTL analyses will uncover additional secondary e/sQTLs, further benefiting GWAS interpretation.

Article Details

Journal Science
Volume / Issue Vol. 393, Issue 6808
Published July 16, 2026
ISSN 0036-8075
Publisher American Association for the Advancement of Science

Journal Info

Science

American Association for the Advancement of Science

ISSN: 0036-8075 Social Sciences

Authors (285)

P

Peter Orchard

Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA.

T

Thomas W. Blackwell

Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.

L

Linda Kachuri

Department of Epidemiology and Population Health, School of Medicine, Stanford University, Stanford, CA, USA.

P

Peter J. Castaldi

Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

M

Michael H. Cho

Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

S

Stephanie A. Christenson

Division of Pulmonary, Critical Care, Allergy, and Sleep Medicine, School of Medicine, University of California, San Francisco, San Francisco, CA, USA.

P

Peter Durda

Department of Pathology and Laboratory Medicine, Larner College of Medicine, University of Vermont, Burlington, VT, USA.

S

Stacey Gabriel

C

Craig P. Hersh

Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

S

Scott Huntsman

Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.

S

Seungyong Hwang

Department of Genetics, Stanford University, Stanford, CA, USA.

R

Roby Joehanes

Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.

M

Mari Johnson

Division of Public Health Sciences, Fred Hutchinson Cancer Center, Seattle, WA, USA.

X

Xingnan Li

Division of Data Driven and Digital Medicine, Department of Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA.

H

Honghuang Lin

Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.

C

Ching-Ti Liu

Department of Biostatistics, School of Public Health, Boston University, Boston, MA, USA.

Y

Yongmei Liu

Division of Cardiology, Department of Medicine, School of Medicine, Duke University, Durham, NC, USA.

A

Angel C. Y. Mak

Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.

A

Ani W. Manichaikul

Department of Genome Sciences, School of Medicine, University of Virginia, Charlottesville, VA, USA.

D

David T. Paik

A

Aabida Saferali

Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

J

Joshua D. Smith

K

Kent D. Taylor

The Institute for Translational Genomics and Population Sciences, Department of Pediatrics, The Lundquist Institute for Biomedical Innovation at Harbor-UCLA Medical Center, Torrance, CA, USA.

R

Russell P. Tracy

Laboratory for Clinical Biochemistry Research, Larner College of Medicine, University of Vermont, Burlington, VT, USA.

J

Jiongming Wang

Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.

M

Mingqiang Wang

J

Joshua S. Weinstock

Department of Biomedical Engineering, Johns Hopkins University, Baltimore, MD, USA.

J

Jeffrey Weiss

H

Heather E. Wheeler

Program in Bioinformatics, Loyola University Chicago, Chicago, IL, USA.

Y

Ying Zhou

S

Sebastian Zöllner

Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.

J

Joseph C. Wu

L

Luisa Mestroni

Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.

S

Sharon Graw

Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.

M

Matthew R. G. Taylor

Department of Medicine, Adult Medical Genetics Program, University of Colorado Anschutz Medical Campus, Aurora, CO, USA.

V

Victor E. Ortega

Department of Medicine, Division of Pulmonary and Critical Care Medicine, Mayo Clinic, Phoenix, AZ, USA.

W

W. Craig Johnson

Collaborative Health Studies Coordinating Center, University of Washington, Seattle, WA, USA.

W

Weiniu Gan

National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.

G

Gonçalo Abecasis

D

Deborah A. Nickerson

Northwest Genomics Center, University of Washington, Seattle, WA, USA.

N

Namrata Gupta

Broad Institute of MIT and Harvard, Cambridge, MA, USA.

K

Kristin Ardlie

Broad Institute of MIT and Harvard, Cambridge, MA, USA.

P

Prescott G. Woodruff

Division of Pulmonary, Critical Care, Allergy, and Sleep Medicine, School of Medicine, University of California, San Francisco, San Francisco, CA, USA.

R

Russell P. Bowler

Department of Genomic Sciences and Systems Biology, Cleveland Clinic, Cleveland, OH, USA.

D

Deborah A. Meyers

Department of Medicine, Mayo Clinic, Scottsdale, AZ, USA.

A

Alex Reiner

Department of Epidemiology, University of Washington, Seattle, WA, USA.

C

Charles Kooperberg

Division of Public Health Sciences, Fred Hutchinson Cancer Center, Seattle, WA, USA.

E

Elad Ziv

Department of Medicine, University of California, San Francisco, San Francisco, CA, USA.

R

Ramachandran S. Vasan

Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.

M

Martin G. Larson

Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.

L

L. Adrienne Cupples

Framingham Heart Study, National Heart, Lung, and Blood Institute, National Institutes of Health and Boston University, Framingham, MA, USA.

E

Edwin K. Silverman

Channing Division of Network Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA.

S

Stephen S. Rich

Department of Genome Sciences, School of Medicine, University of Virginia, Charlottesville, VA, USA.

N

Nancy Heard-Costa

Chobanian & Avedisian School of Medicine, Boston University, Boston, MA, USA.

H

Hua Tang

Department of Genetics, Stanford University, Stanford, CA, USA.

J

Jerome I. Rotter

A

Albert V. Smith

Department of Biostatistics, School of Public Health, University of Michigan, Ann Arbor, MI, USA.

D

Daniel Levy

Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, MD, USA.

F

François Aguet

L

Laura J. Scott

L

Laura M. Raffield

Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.

S

Stephen C. J. Parker

Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA.

N

Namiko Abe

New York Genome Center, New York, NY, USA.

L

Laura Almasy

Children’s Hospital of Philadelphia, University of Pennsylvania, Philadelphia, PA, USA.

S

Seth Ament

University of Maryland, Baltimore, MD, USA.

P

Pramod Anugu

University of Mississippi, Jackson, MS, USA.

P

Paul Auer

Medical College of Wisconsin, Milwaukee, WI, USA.

D

Dimitrios Avramopoulos

Johns Hopkins University, Baltimore, MD, USA.

A

Adithya Balasubramanian

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

R

R. Graham Barr

Columbia University, New York, NY, USA.

L

Lucas Barwick

The Emmes Corporation, Rockville, MD, USA.

T

Terri Beaty

Johns Hopkins University, Baltimore, MD, USA.

D

Diane Becker

Johns Hopkins University, Baltimore, MD, USA.

L

Lewis Becker

Johns Hopkins University, Baltimore, MD, USA.

A

Amber Beitelshees

University of Maryland, Baltimore, MD, USA.

T

Takis Benos

University of Pittsburgh, Pittsburgh, PA, USA.

M

Marcos Bezerra

Fundação de Hematologia e Hemoterapia de Pernambuco - Hemope, Recife, Brazil.

J

Joshua Bis

University of Washington, Seattle, WA, USA.

J

Jennifer Brody

University of Washington, Seattle, WA, USA.

U

Ulrich Broeckel

Medical College of Wisconsin, Milwaukee, WI, USA.

J

Jai Broome

University of Washington, Seattle, WA, USA.

K

Karen Bunting

New York Genome Center, New York, NY, USA.

E

Erin Buth

University of Washington, Seattle, WA, USA.

V

Vincent Carey

Brigham andWomen’s Hospital, Boston, MA, USA.

C

Cara Carty

Washington State University, Pullman, WA, USA.

R

Richard Casaburi

University of California, Los Angeles, Los Angeles, CA, USA.

M

Mark Chaffin

Broad Institute, Cambridge, MA, USA.

C

Christy Chang

University of Maryland, Baltimore, MD, USA.

Y

Yi-Cheng Chang

National Taiwan University, Taipei, Taiwan (Province of China).

S

Sameer Chavan

University of Colorado at Denver, Denver, CO, USA.

B

Bo-Juen Chen

New York Genome Center, New York, NY, USA.

W

Wei-Min Chen

University of Virginia, Charlottesville, VA, USA.

S

Seung Hoan Choi

Broad Institute, Cambridge, MA, USA.

L

Lee-Ming Chuang

National Taiwan University, Taipei, Taiwan (Province of China).

R

Ren-Hua Chung

National Health Research Institute Taiwan, Miaoli County, Taiwan (Province of China).

M

Matthew Conomos

University of Washington, Seattle, WA, USA.

E

Elaine Cornell

University of Vermont, Burlington, VT, USA.

C

Carolyn Crandall

University of California, Los Angeles, Los Angeles, CA, USA.

J

James Crapo

National Jewish Health, Denver, CO, USA.

J

Jeffrey Curtis

University of Michigan, Ann Arbor, MI, USA.

C

Coleen Damcott

University of Maryland, Baltimore, MD, USA.

S

Sean David

University of Chicago, Chicago, IL, USA.

L

Lisa de las Fuentes

Washington University in St. Louis, St. Louis, MO, USA.

P

Paul de Vries

University of Texas Health at Houston, Houston, TX, USA.

R

Ranjan Deka

University of Cincinnati, Cincinnati, OH, USA.

D

Dawn DeMeo

Brigham andWomen’s Hospital, Boston, MA, USA.

S

Scott Devine

University of Maryland, Baltimore, MD, USA.

H

Huyen Dinh

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

H

Harsha Doddapaneni

Q

Qing Duan

University of North Carolina, Chapel Hill, NC, USA.

R

Ravi Duggirala

University of Texas Rio Grande Valley School of Medicine, Edinburg, TX, USA.

C

Charles Eaton

Harvard University, Cambridge, MA, USA.

L

Lynette Ekunwe

University of Mississippi, Jackson, MS, USA.

A

Adel El Boueiz

Washington University in St. Louis, St. Louis, MO, USA.

L

Leslie Emery

University of Washington, Seattle, WA, USA.

C

Charles Farber

University of Virginia, Charlottesville, VA, USA.

J

Jesse Farek

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

N

Nora Franceschini

University of North Carolina, Chapel Hill, NC, USA.

C

Chris Frazar

University of Washington, Seattle, WA, USA.

M

Mao Fu

University of Maryland, Baltimore, MD, USA.

S

Stephanie M. Fullerton

University of Washington, Seattle, WA, USA.

L

Lucinda Fulton

Washington University in St. Louis, St. Louis, MO, USA.

S

Shanshan Gao

University of Colorado at Denver, Denver, CO, USA.

Y

Yan Gao

M

Margery Gass

Fred Hutchinson Cancer Research Center, Seattle, WA, USA.

H

Heather Geiger

A

Auyon Ghosh

Brigham andWomen’s Hospital, Boston, MA, USA.

C

Chris Gignoux

Stanford University, Stanford, CA, USA.

D

David Glahn

Boston Children’s Hospital, Harvard Medical School, Boston, MA, USA.

S

Stephanie Gogarten

University of Washington, Seattle, WA, USA.

D

Da-Wei Gong

University of Maryland, Baltimore, MD, USA.

H

Harald Goring

University of Texas Rio Grande Valley School of Medicine, San Antonio, TX, USA.

D

Daniel Grine

University of Colorado at Denver, Denver, CO, USA.

C

C. Charles Gu

Washington University in St. Louis, St. Louis, MO, USA.

Y

Yue Guan

University of Maryland, Baltimore, MD, USA.

M

Michael Hall

University of Mississippi, Jackson, MS, USA.

Y

Yi Han

D

Daniel Harris

University of Maryland, Philadelphia, PA, USA.

B

Ben Heavner

D

David Herrington

Wake Forest Baptist Health, Winston-Salem, NC, USA.

B

Brian Hobbs

E

Elliott Hong

University of Maryland, Baltimore, MD, USA.

K

Karin Hoth

University of Iowa, Iowa City, IA.

C

Chao (Agnes) Hsiung

National Health Research Institute Taiwan, Miaoli County, Taiwan (Province of China).

J

Jianhong Hu

Y

Yi-Jen Hung

Tri-Service General Hospital National Defense Medical Center, Taiwan (Province of China).

H

Haley Huston

Blood Works Northwest, Seattle, WA, USA.

C

Chii Min Hwu

Taichung Veterans General Hospital Taiwan, Taichung City, Taiwan (Province of China).

R

Rebecca Jackson

D

Deepti Jain

University of Washington, Seattle, WA, USA.

J

Jill Johnsen

University of Washington, Seattle, WA, USA.

R

Rich Johnston

Emory University, Atlanta, GA, USA.

K

Kimberly Jones

Johns Hopkins University, Baltimore, MD, USA.

M

Michael Kessler

A

Alyna Khan

University of Washington, Seattle, WA, USA.

Z

Ziad Khan

W

Wonji Kim

Harvard University, Cambridge, MA, USA.

J

John Kimoff

McGill University, Montréal, QC, Canada.

G

Greg Kinney

University of Colorado at Denver, Aurora, CO, USA.

H

Holly Kramer

Loyola University, Maywood, IL, USA.

C

Christoph Lange

Harvard School of Public Health, Boston, MA, USA.

E

Ethan Lange

University of Colorado at Denver, Denver, CO, USA.

C

Cathy Laurie

University of Washington, Seattle, WA, USA.

C

Cecelia Laurie

University of Washington, Seattle, WA, USA.

M

Meryl LeBoff

Brigham andWomen’s Hospital, Boston, MA, USA.

S

Sandra Lee

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

W

Wen-Jane Lee

Taichung Veterans General Hospital Taiwan, Taichung City, Taiwan (Province of China).

D

David Levine

University of Washington, Seattle, WA, USA.

J

Joshua Lewis

University of Maryland, Baltimore, MD, USA.

Y

Yun Li

X

Xihong Lin

S

Simin Liu

Brown University, Providence, RI, USA.

Y

Yu Liu

B

Barry Make

Johns Hopkins University, Baltimore, MD, USA.

A

Alisa Manning

Broad Institute, Harvard University, Massachusetts General Hospital, Boston, MA, USA.

J

JoAnn Manson

Brigham andWomen’s Hospital, Boston, MA, USA.

L

Lisa Martin

M

Melissa Marton

New York Genome Center, New York City, NY, USA.

S

Susan Mathai

University of Colorado at Denver, Denver, CO, USA.

S

Susanne May

P

Patrick McArdle

University of Maryland, Baltimore, MD, USA.

M

Merry-Lynn McDonald

S

Sean McFarland

Harvard University, Cambridge, MA, USA.

D

Daniel McGoldrick

University of Washington, Seattle, WA, USA.

C

Caitlin McHugh

University of Washington, Seattle, WA, USA.

H

Hao Mei

University of Mississippi, Jackson, MS, USA.

J

James Meigs

Massachusetts General Hospital, Boston, MA, USA.

V

Vipin Menon

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

N

Nancy Min

University of Mississippi, Jackson, MS, USA.

M

Matt Moll

Brigham andWomen’s Hospital, Boston, MA, USA.

Z

Zeineen Momin

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

M

May Montasser

National Heart, Lung, and Blood Institute, Bethesda, MD, USA.

J

Josyf C Mychaleckyj

University of Virginia, Charlottesville, VA, USA.

R

Rakhi Naik

Johns Hopkins University, Baltimore, MD, USA.

T

Take Naseri

P

Pradeep Natarajan

S

Sarah C. Nelson

B

Bonnie Neltner

University of Colorado at Denver, Denver, CO, USA.

C

Caitlin Nessner

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

O

Osuji Nkechinyere

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

J

Jeff O'Connell

University of Maryland, Baltimore, MD, USA.

T

Tim O’Connor

University of Maryland, Baltimore, MD, USA.

H

Heather Ochs-Balcom

University at Buffalo, Buffalo, NY, USA.

G

Geoffrey Okwuonu

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

J

James Pankow

University of Minnesota, Minneapolis, MN, USA.

C

Cora Parker

RTI International, Research Triangle Park, NC, USA.

G

Gina Peloso

Boston University, Boston, MA, USA.

J

Juan Manuel Peralta

University of Texas Rio Grande Valley School of Medicine, Edinburg, TX, USA.

M

Marco Perez

Stanford University, Stanford, CA, USA.

J

James Perry

University of Maryland, Baltimore, MD, USA.

U

Ulrike Peters

Fred Hutchinson Cancer Research Center, Seattle, WA, USA.

L

Lawrence S Phillips

Emory University, Atlanta, GA, USA.

T

Toni Pollin

University of Maryland, Baltimore, MD, USA.

J

Julia Powers Becker

University of Colorado at Denver, Denver, CO, USA.

M

Meher Preethi Boorgula

University of Colorado at Denver, Denver, CO, USA.

B

Bruce Psaty

University of Washington, Seattle, WA, USA.

D

Dandi Qiao

Brigham andWomen’s Hospital, Boston, MA, USA.

N

Nicholas Rafaels

University of Colorado at Denver, Denver, CO, USA.

M

Mahitha Rajendran

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

L

Laura Rasmussen-Torvik

Northwestern University, Chicago, IL, USA.

A

Aakrosh Ratan

University of Virginia, Charlottesville, VA, USA.

R

Robert Reed

University of Maryland, Baltimore, MD, USA.

E

Elizabeth Regan

National Jewish Health, Denver, CO, USA.

M

Muagututi‘a Sefuiva Reupena

Lutia I Puava Ae Mapu I Fagalele, Apia, Samoa.

R

Rebecca Robillard

University of Ottawa, Ottawa, ON, Canada.

C

Carolina Roselli

Broad Institute, Cambridge, MA, USA.

I

Ingo Ruczinski

Johns Hopkins University, Baltimore, MD, USA.

A

Alexi Runnels

P

Pamela Russell

University of Colorado at Denver, Denver, CO, USA.

K

Kathleen Ryan

University of Maryland, Baltimore, MD, USA.

E

Ester Cerdeira Sabino

Universidade de Sao Paulo, Sao Paulo, Brazil.

S

Shabnam Salimi

University of Maryland, Seattle, WA, USA.

S

Sejal Salvi

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

S

Steven Salzberg

Johns Hopkins University, Baltimore, MD, USA.

K

Kevin Sandow

Lundquist Institute, Torrance, CA, USA.

J

Jireh Santibanez

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

K

Karen Schwander

Washington University in St. Louis, St. Louis, MO, USA.

F

Frank Sciurba

University of Pittsburgh, Pittsburgh, PA, USA.

F

Frédéric Sériès

Université Laval, Quebec City, Canada.

A

Amol Shetty

University of Maryland, Baltimore, MD, USA.

A

Aniket Shetty

University of Colorado at Denver, Denver, CO, USA.

B

Brian Silver

UMass Memorial Medical Center, Worcester, MA, USA.

R

Robert Skomro

University of Saskatchewan, Saskatoon, Canada.

T

Tanja Smith

New York Genome Center, New York, NY, USA.

S

Sylvia Smoller

Albert Einstein College of Medicine, New York, NY, USA.

B

Beverly Snively

Wake Forest Baptist Health, Winston-Salem, NC, USA.

A

Adrienne M. Stilp

G

Garrett Storm

University of Colorado at Denver, Aurora, CO, USA.

E

Elizabeth Streeten

University of Maryland, Baltimore, MD, USA.

J

Jessica Lasky Su

Brigham andWomen’s Hospital, Boston, MA, USA.

Y

Yun Ju Sung

Washington University in St. Louis, St. Louis, MO, USA.

J

Jody Sylvia

Brigham andWomen’s Hospital, Boston, MA, USA.

A

Adam Szpiro

University of Washington, Seattle, WA, USA.

M

Margaret Taub

Johns Hopkins University, Baltimore, MD, USA.

S

Simeon Taylor

University of Maryland, Baltimore, MD, USA.

T

Timothy A. Thornton

University of Washington, Seattle, WA, USA.

M

Machiko Threlkeld

University of Washington, Seattle, WA, USA.

L

Lesley Tinker

Fred Hutchinson Cancer Research Center, Seattle, WA, USA.

D

David Tirschwell

University of Washington, Seattle, WA, USA.

H

Hemant Tiwari

University of Alabama, Birmingham, AL, USA.

C

Catherine Tong

University of Washington, Seattle, WA, USA.

M

Michael Tsai

University of Minnesota, Minneapolis, MN, USA.

D

Dhananjay Vaidya

Johns Hopkins University, Baltimore, MD, USA.

T

Tarik Walker

University of Colorado at Denver, Denver, CO, USA.

R

Robert Wallace

University of Iowa, Iowa City, IA.

A

Avram Walts

University of Colorado at Denver, Denver, CO, USA.

F

Fei Fei Wang

University of Washington, Seattle, WA, USA.

H

Heming Wang

K

Karol Watson

University of California, Los Angeles, Los Angeles, CA, USA.

J

Jennifer Watt

Baylor College of Medicine Human Genome Sequencing Center, Houston, TX, USA.

L

Lu-Chen Weng

Massachusetts General Hospital, Boston, MA, USA.

J

Jennifer Wessel

Indiana University, Indianapolis, IN, USA.

K

Kayleen Williams

University of Washington, Seattle, WA, USA.

C

Carla Wilson

Brigham andWomen’s Hospital, Boston, MA, USA.

J

James Wilson

L

Lara Winterkorn

New York Genome Center, New York City, NY, USA.

Q

Quenna Wong

B

Baojun Wu

Henry Ford Health System, Detroit, MI, USA.

H

Huichun Xu

University of Maryland, Baltimore, MD, USA.

L

Lisa Yanek

Johns Hopkins University, Baltimore, MD, USA.

I

Ivana Yang

University of Colorado at Denver, Denver, CO, USA.

S

Seyedeh Maryam Zekavat

Broad Institute, Cambridge, MA, USA.

S

Snow Xueyan Zhao

National Jewish Health, Denver, CO, USA.

W

Wei Zhao

X

Xiaofeng Zhu