title: Exome Genotyping Identifies Pleiotropic Variants Associated with Red Blood Cell Traits.
abstract: Red blood cell (RBC) traits are important heritable clinical biomarkers and modifiers of disease severity. To identify coding genetic variants associated with these traits, we conducted meta-analyses of seven RBC phenotypes in 130,273 multi-ethnic individuals from studies genotyped on an exome array. After conditional analyses and replication in 27,480 independent individuals, we identified 16 new RBC variants. We found low-frequency missense variants in MAP1A (rs55707100, minor allele frequency [MAF] = 3.3%, p = 2 × 10(-10) for hemoglobin [HGB]) and HNF4A (rs1800961, MAF = 2.4%, p < 3 × 10(-8) for hematocrit [HCT] and HGB). In African Americans, we identified a nonsense variant in CD36 associated with higher RBC distribution width (rs3211938, MAF = 8.7%, p = 7 × 10(-11)) and showed that it is associated with lower CD36 expression and strong allelic imbalance in ex vivo differentiated human erythroblasts. We also identified a rare missense variant in ALAS2 (rs201062903, MAF = 0.2%) associated with lower mean corpuscular volume and mean corpuscular hemoglobin (p < 8 × 10(-9)). Mendelian mutations in ALAS2 are a cause of sideroblastic anemia and erythropoietic protoporphyria. Gene-based testing highlighted three rare missense variants in PKLR, a gene mutated in Mendelian non-spherocytic hemolytic anemia, associated with HGB and HCT (SKAT p < 8 × 10(-7)). These rare, low-frequency, and common RBC variants showed pleiotropy, being also associated with platelet, white blood cell, and lipid traits. Our association results and functional annotation suggest the involvement of new genes in human erythropoiesis. We also confirm that rare and low-frequency variants play a role in the architecture of complex human traits, although their phenotypic effect is generally smaller than originally anticipated.
authors.affiliation: Department of Medicine, Université de Montréal, Montréal, QC H3T 1J4, Canada; Montreal Heart Institute, Montréal, QC H1T 1C8, Canada.
authors.forename: Nathalie
authors.initials: N
authors.lastname: Chami
authors.affiliation: Population Sciences Branch, National Heart, Lung, and Blood Institute, The Framingham Heart Study, Framingham, MA 01702, USA.
authors.forename: Ming-Huei
authors.initials: MH
authors.lastname: Chen
authors.affiliation: Genetics Target Sciences, GlaxoSmithKline, Research Triangle Park, NC 27709, USA; OmicSoft Corporation, Cary, NC 27513, USA.
authors.forename: Andrew J
authors.initials: AJ
authors.lastname: Slater
authors.affiliation: Population Sciences Branch, National Heart, Lung, and Blood Institute, The Framingham Heart Study, Framingham, MA 01702, USA.
authors.forename: John D
authors.initials: JD
authors.lastname: Eicher
authors.affiliation: Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK; Department of Hygiene and Epidemiology, University of Ioannina Medical School, Ioannina 45110, Greece.
authors.forename: Evangelos
authors.initials: E
authors.lastname: Evangelou
authors.affiliation: Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Baltimore, MD 21224, USA.
authors.forename: Salman M
authors.initials: SM
authors.lastname: Tajuddin
authors.affiliation: Department of Medicine, Center of Human Nutrition, Washington University School of Medicine, St Louis, MO 63110, USA.
authors.forename: Latisha
authors.initials: L
authors.lastname: Love-Gregory
authors.affiliation: Department of Functional Genomics, Interfaculty Institute for Genetics and Functional Genomics, University Medicine, Greifswald and Ernst-Mortiz-Arndt University Greifswald, Greifswald 17475, Germany; DZHK (German Centre for Cardiovascular Research), partner site Greifswald, Greifswald QA, Germany.
authors.forename: Tim
authors.initials: T
authors.lastname: Kacprowski
authors.affiliation: The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA.
authors.forename: Ursula M
authors.initials: UM
authors.lastname: Schick
authors.affiliation: Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA 02114, USA; Program in Medical and Population Genetics, Broad Institute, Cambridge, MA 02142, USA; Cardiovascular Research Center, Massachusetts General Hospital, Boston, MA 02114, USA; Department of Medicine, Harvard Medical School, Boston, MA 02115, USA; Division of Cardiovascular Medicine, Kanazawa University, Graduate School of Medical Science, Kanazawa, Ishikawa 9200942, Japan.
authors.forename: Akihiro
authors.initials: A
authors.lastname: Nomura
authors.affiliation: Division of Epidemiology, Department of Medicine, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, TN 37235, USA.
authors.forename: Ayush
authors.initials: A
authors.lastname: Giri
authors.affiliation: Department of Medicine, Université de Montréal, Montréal, QC H3T 1J4, Canada; Montreal Heart Institute, Montréal, QC H1T 1C8, Canada.
authors.forename: Samuel
authors.initials: S
authors.lastname: Lessard
authors.affiliation: Department of Medicine, University of Washington, Seattle, WA 98101, USA.
authors.forename: Jennifer A
authors.initials: JA
authors.lastname: Brody
authors.affiliation: The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA; The Genetics of Obesity and Related Metabolic Traits Program, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA.
authors.forename: Claudia
authors.initials: C
authors.lastname: Schurmann
authors.affiliation: Department of Laboratory Medicine and Pathology, University of Minnesota, Minneapolis, MN 55454, USA.
authors.forename: Nathan
authors.initials: N
authors.lastname: Pankratz
authors.affiliation: Department of Medicine/Division of General Internal Medicine, Johns Hopkins University, School of Medicine, Baltimore, MD 21205, USA.
authors.forename: Lisa R
authors.initials: LR
authors.lastname: Yanek
authors.affiliation: Center for Public Health Genomics, University of Virginia, Charlottesville, VA 22908, USA.
authors.forename: Ani
authors.initials: A
authors.lastname: Manichaikul
authors.affiliation: Department of Epidemiology, Erasmus, MC Rotterdam 3000, the Netherlands.
authors.forename: Raha
authors.initials: R
authors.lastname: Pazoki
authors.affiliation: Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.
authors.forename: Evelin
authors.initials: E
authors.lastname: Mihailov
authors.affiliation: Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh EH8 9JZ, UK; Department of Psychology, University of Edinburgh, Edinburgh EH8 9JZ, UK.
authors.forename: W David
authors.initials: WD
authors.lastname: Hill
authors.affiliation: Department of Genetics, University of North Carolina, Chapel Hill, NC 27514, USA.
authors.forename: Laura M
authors.initials: LM
authors.lastname: Raffield
authors.affiliation: Division of Medical Genetics, Department of Medicine, University of Washington, Seattle, WA 98195, USA.
authors.forename: Amber
authors.initials: A
authors.lastname: Burt
authors.affiliation: Department of Biostatistics, University of Washington, Seattle, WA 98195, USA.
authors.forename: Traci M
authors.initials: TM
authors.lastname: Bartz
authors.affiliation: Department of Medicine/Division of General Internal Medicine, Johns Hopkins University, School of Medicine, Baltimore, MD 21205, USA.
authors.forename: Diane M
authors.initials: DM
authors.lastname: Becker
authors.affiliation: Department of Medicine/Divisions of Cardiology and General Internal Medicine, Johns Hopkins University School of Medicine, Baltimore, MD 21205, USA.
authors.forename: Lewis C
authors.initials: LC
authors.lastname: Becker
authors.affiliation: Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX 77030, USA; Human Genome Sequencing Center, Baylor College of Medicine, Houston, TX 77030, USA.
authors.forename: Eric
authors.initials: E
authors.lastname: Boerwinkle
authors.affiliation: The Novo Nordisk Foundation, Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen 2100, Denmark.
authors.forename: Jette
authors.initials: J
authors.lastname: Bork-Jensen
authors.affiliation: The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA.
authors.forename: Erwin P
authors.initials: EP
authors.lastname: Bottinger
authors.affiliation: TIMI Study Group, Cardiovascular Division, Brigham and Women's Hospital, Boston, MA 02115, USA.
authors.forename: Michelle L
authors.initials: ML
authors.lastname: O'Donoghue
authors.affiliation: Department of Biomedical Informatics and Medical Education, University of Washington, Seattle, WA 98195, USA.
authors.forename: David R
authors.initials: DR
authors.lastname: Crosslin
authors.affiliation: Montreal Heart Institute, Montréal, QC H1T 1C8, Canada; Faculty of Pharmacy, Université de Montréal, Montréal, QC H3T 1J4, Canada.
authors.forename: Simon
authors.initials: S
authors.lastname: de Denus
authors.affiliation: Department of Medicine, Université de Montréal, Montréal, QC H3T 1J4, Canada; Montreal Heart Institute, Montréal, QC H1T 1C8, Canada.
authors.forename: Marie-Pierre
authors.initials: MP
authors.lastname: Dubé
authors.affiliation: Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.
authors.forename: Paul
authors.initials: P
authors.lastname: Elliott
authors.affiliation: Department of Clinical Sciences, Malmö, Lund University, Malmö 221 00, Sweden; Skåne University Hospital, Malmö 222 41, Sweden.
authors.forename: Gunnar
authors.initials: G
authors.lastname: Engström
authors.affiliation: Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Baltimore, MD 21224, USA.
authors.forename: Michele K
authors.initials: MK
authors.lastname: Evans
authors.affiliation: Department of Medicine, University of Washington, Seattle, WA 98101, USA.
authors.forename: James S
authors.initials: JS
authors.lastname: Floyd
authors.affiliation: Institute of Molecular Medicine, The University of Texas Health Science Center at Houston, Houston, TX 77030, USA.
authors.forename: Myriam
authors.initials: M
authors.lastname: Fornage
authors.affiliation: Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.
authors.forename: He
authors.initials: H
authors.lastname: Gao
authors.affiliation: Institute for Immunology and Transfusion Medicine, University Medicine Greifswald, Greifswald 17475, Germany.
authors.forename: Andreas
authors.initials: A
authors.lastname: Greinacher
authors.affiliation: Icelandic Heart Association, 201 Kopavogur, Iceland; Faculty of Medicine, University of Iceland, 101 Reykjavik, Iceland.
authors.forename: Vilmundur
authors.initials: V
authors.lastname: Gudnason
authors.affiliation: The Novo Nordisk Foundation, Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen 2100, Denmark.
authors.forename: Torben
authors.initials: T
authors.lastname: Hansen
authors.affiliation: Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, Intramural Research Program, NIH, Bethesda, MD 20892, USA.
authors.forename: Tamara B
authors.initials: TB
authors.lastname: Harris
authors.affiliation: MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh EH4 2XU, UK.
authors.forename: Caroline
authors.initials: C
authors.lastname: Hayward
authors.affiliation: Department of Clinical Chemistry, Fimlab Laboratories, Tampere 33520, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere 33014, Finland; University of Tampere, School of Medicine, Tampere 33014, Finland.
authors.forename: Jussi
authors.initials: J
authors.lastname: Hernesniemi
authors.affiliation: Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX 77030, USA; Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, NC 27514, USA.
authors.forename: Heather M
authors.initials: HM
authors.lastname: Highland
authors.affiliation: Program in Medical and Population Genetics, Broad Institute, Cambridge, MA 02142, USA; Department of Endocrinology, Boston Children's Hospital, Boston, MA 02115, USA.
authors.forename: Joel N
authors.initials: JN
authors.lastname: Hirschhorn
authors.affiliation: Department of Epidemiology, Erasmus, MC Rotterdam 3000, the Netherlands; Department of Epidemiology, Harvard TH Chan School of Public Health, Boston, MA 02115, USA.
authors.forename: Albert
authors.initials: A
authors.lastname: Hofman
authors.affiliation: Department of Epidemiology, School of Public Health, University of Alabama at Birmingham, Birmingham, AL 35233, USA.
authors.forename: Marguerite R
authors.initials: MR
authors.lastname: Irvin
authors.affiliation: Department of Clinical Physiology, Tampere University Hospital, Tampere 33521, Finland; Department of Clinical Physiology, University of Tampere School of Medicine, Tampere 33014, Finland.
authors.forename: Mika
authors.initials: M
authors.lastname: Kähönen
authors.affiliation: Departments of Genetics and Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, NC 27599, USA.
authors.forename: Ethan
authors.initials: E
authors.lastname: Lange
authors.affiliation: Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, Intramural Research Program, NIH, Bethesda, MD 20892, USA.
authors.forename: Lenore J
authors.initials: LJ
authors.lastname: Launer
authors.affiliation: Department of Clinical Chemistry, Fimlab Laboratories, Tampere 33520, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere 33014, Finland.
authors.forename: Terho
authors.initials: T
authors.lastname: Lehtimäki
authors.affiliation: Department of Medicine, Division of Cardiovascular Medicine, Stanford University, School of Medicine, Palo Alto, CA 94305, USA.
authors.forename: Jin
authors.initials: J
authors.lastname: Li
authors.affiliation: Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh EH8 9JZ, UK; Department of Psychology, University of Edinburgh, Edinburgh EH8 9JZ, UK.
authors.forename: David C M
authors.initials: DC
authors.lastname: Liewald
authors.affiliation: Research Centre for Prevention and Health, The Capital Region of Denmark, Copenhagen 2600, Denmark; Department of Clinical Experimental Research, Rigshospitalet, Glostrup 2100, Denmark; Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen 2200, Denmark.
authors.forename: Allan
authors.initials: A
authors.lastname: Linneberg
authors.affiliation: Center for Human Genetics, Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, NC 27157, USA.
authors.forename: Yongmei
authors.initials: Y
authors.lastname: Liu
authors.affiliation: The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA; The Genetics of Obesity and Related Metabolic Traits Program, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA.
authors.forename: Yingchang
authors.initials: Y
authors.lastname: Lu
authors.affiliation: Department of Clinical Chemistry, Fimlab Laboratories, Tampere 33520, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere 33014, Finland.
authors.forename: Leo-Pekka
authors.initials: LP
authors.lastname: Lyytikäinen
authors.affiliation: Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.
authors.forename: Reedik
authors.initials: R
authors.lastname: Mägi
authors.affiliation: Department of Medicine, Divisions of Allergy and Clinical Immunology and General Internal Medicine, Johns Hopkins University School of Medicine, Baltimore, MD 21205, USA.
authors.forename: Rasika A
authors.initials: RA
authors.lastname: Mathias
authors.affiliation: Department of Clinical Sciences, Malmö, Lund University, Malmö 221 00, Sweden; Skåne University Hospital, Malmö 222 41, Sweden.
authors.forename: Olle
authors.initials: O
authors.lastname: Melander
authors.affiliation: Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.
authors.forename: Andres
authors.initials: A
authors.lastname: Metspalu
authors.affiliation: Department of Clinical Chemistry, Fimlab Laboratories, Tampere 33520, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere 33014, Finland.
authors.forename: Nina
authors.initials: N
authors.lastname: Mononen
authors.affiliation: Laboratory of Neurogenetics, National Institute on Aging, NIH, Bethesda, MD 20892, USA.
authors.forename: Mike A
authors.initials: MA
authors.lastname: Nalls
authors.affiliation: Department of Genome Sciences, University of Washington, Seattle, WA 98105, USA.
authors.forename: Deborah A
authors.initials: DA
authors.lastname: Nickerson
authors.affiliation: University of Tampere, School of Medicine, Tampere 33014, Finland; Department of Cardiology, Heart Center, Tampere University Hospital, Tampere 33521, Finland.
authors.forename: Kjell
authors.initials: K
authors.lastname: Nikus
authors.affiliation: Population Sciences Branch, National Heart, Lung, and Blood Institute, The Framingham Heart Study, Framingham, MA 01702, USA; Cardiology Section and Center for Population Genomics, Boston Veteran's Administration (VA) Healthcare, Boston, MA 02118, USA.
authors.forename: Chris J
authors.initials: CJ
authors.lastname: O'Donnell
authors.affiliation: Department of Clinical Sciences, Malmö, Lund University, Malmö 221 00, Sweden; Skåne University Hospital, Malmö 222 41, Sweden.
authors.forename: Marju
authors.initials: M
authors.lastname: Orho-Melander
authors.affiliation: The Novo Nordisk Foundation, Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen 2100, Denmark.
authors.forename: Oluf
authors.initials: O
authors.lastname: Pedersen
authors.affiliation: Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald 17475, Germany.
authors.forename: Astrid
authors.initials: A
authors.lastname: Petersmann
authors.affiliation: Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX 77030, USA.
authors.forename: Linda
authors.initials: L
authors.lastname: Polfus
authors.affiliation: Cardiovascular Health Research Unit, Departments of Medicine Epidemiology and Health Services, University of Washington, Seattle, WA 98101, USA; Group Health Research Institute, Group Health Cooperative, Seattle, WA 98101, USA.
authors.forename: Bruce M
authors.initials: BM
authors.lastname: Psaty
authors.affiliation: Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku 20521, Finland; Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku 20520, Finland.
authors.forename: Olli T
authors.initials: OT
authors.lastname: Raitakari
authors.affiliation: Department of Clinical Chemistry, Fimlab Laboratories, Tampere 33520, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere 33014, Finland.
authors.forename: Emma
authors.initials: E
authors.lastname: Raitoharju
authors.affiliation: Institute of Molecular Medicine, The University of Texas Health Science Center at Houston, Houston, TX 77030, USA.
authors.forename: Melissa
authors.initials: M
authors.lastname: Richard
authors.affiliation: Department of Biostatistics, University of Washington, Seattle, WA 98195, USA.
authors.forename: Kenneth M
authors.initials: KM
authors.lastname: Rice
authors.affiliation: Department of Epidemiology, Erasmus, MC Rotterdam 3000, the Netherlands; Department of Internal Medicine, Erasmus MC, Rotterdam 3000, the Netherlands; Netherlands Consortium for Healthy Ageing (NCHA), Rotterdam 3015, the Netherlands.
authors.forename: Fernando
authors.initials: F
authors.lastname: Rivadeneira
authors.affiliation: Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute, Torrance, CA 90502, USA; Department of Pediatrics, Harbor-UCLA Medical Center, Torrance, CA 90502, USA.
authors.forename: Jerome I
authors.initials: JI
authors.lastname: Rotter
authors.affiliation: Department of Functional Genomics, Interfaculty Institute for Genetics and Functional Genomics, University Medicine, Greifswald and Ernst-Mortiz-Arndt University Greifswald, Greifswald 17475, Germany.
authors.forename: Frank
authors.initials: F
authors.lastname: Schmidt
authors.affiliation: Icelandic Heart Association, 201 Kopavogur, Iceland; Faculty of Medicine, University of Iceland, 101 Reykjavik, Iceland.
authors.forename: Albert Vernon
authors.initials: AV
authors.lastname: Smith
authors.affiliation: Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh EH8 9JZ, UK; Alzheimer Scotland Research Centre, Edinburgh EH8 9JZ, UK.
authors.forename: John M
authors.initials: JM
authors.lastname: Starr
authors.affiliation: Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute, Torrance, CA 90502, USA; Department of Pediatrics, Harbor-UCLA Medical Center, Torrance, CA 90502, USA.
authors.forename: Kent D
authors.initials: KD
authors.lastname: Taylor
authors.affiliation: Institute for Community Medicine, University Medicine Greifswald, Greifswald 17475, Germany.
authors.forename: Alexander
authors.initials: A
authors.lastname: Teumer
authors.affiliation: Research Centre for Prevention and Health, The Capital Region of Denmark, Copenhagen 2600, Denmark.
authors.forename: Betina H
authors.initials: BH
authors.lastname: Thuesen
authors.affiliation: Division of Epidemiology, Department of Medicine, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, TN 37235, USA.
authors.forename: Eric S
authors.initials: ES
authors.lastname: Torstenson
authors.affiliation: Departments of Pathology and Laboratory Medicine and Biochemistry, University of Vermont College of Medicine, Colchester, VT 05446, USA.
authors.forename: Russell P
authors.initials: RP
authors.lastname: Tracy
authors.affiliation: Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK; Department of Hygiene and Epidemiology, University of Ioannina Medical School, Ioannina 45110, Greece.
authors.forename: Ioanna
authors.initials: I
authors.lastname: Tzoulaki
authors.affiliation: Departments of Medicine and Pathology, University of Vermont College of Medicine, Burlington, VT 05405, USA.
authors.forename: Neil A
authors.initials: NA
authors.lastname: Zakai
authors.affiliation: Department of Family Population and Preventive Medicine, Stony Brook University, Stony Brook, NY 11794, USA.
authors.forename: Caterina
authors.initials: C
authors.lastname: Vacchi-Suzzi
authors.affiliation: Department of Epidemiology, Erasmus, MC Rotterdam 3000, the Netherlands.
authors.forename: Cornelia M
authors.initials: CM
authors.lastname: van Duijn
authors.affiliation: Department of Epidemiology, Erasmus, MC Rotterdam 3000, the Netherlands.
authors.forename: Frank J A
authors.initials: FJ
authors.lastname: van Rooij
authors.affiliation: Departments of Medicine and Pathology, University of Vermont College of Medicine, Burlington, VT 05405, USA.
authors.forename: Mary
authors.initials: M
authors.lastname: Cushman
authors.affiliation: Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh EH8 9JZ, UK; Department of Psychology, University of Edinburgh, Edinburgh EH8 9JZ, UK.
authors.forename: Ian J
authors.initials: IJ
authors.lastname: Deary
authors.affiliation: Vanderbilt Epidemiology Center, Department of Obstetrics & Gynecology, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, TN 37203, USA.
authors.forename: Digna R
authors.initials: DR
authors.lastname: Velez Edwards
authors.affiliation: Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.
authors.forename: Anne-Claire
authors.initials: AC
authors.lastname: Vergnaud
authors.affiliation: Department of Medical Sciences, Cardiology and Uppsala Clinical Research Center, Uppsala University, Uppsala 751 85, Sweden.
authors.forename: Lars
authors.initials: L
authors.lastname: Wallentin
authors.affiliation: Genetics Target Sciences, GlaxoSmithKline, King of Prussia, PA 19406, USA.
authors.forename: Dawn M
authors.initials: DM
authors.lastname: Waterworth
authors.affiliation: Green Lane Cardiovascular Service, Auckland City Hospital and University of Auckland, Auckland 1142, New Zealand.
authors.forename: Harvey D
authors.initials: HD
authors.lastname: White
authors.affiliation: Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, MS 39216, USA.
authors.forename: James G
authors.initials: JG
authors.lastname: Wilson
authors.affiliation: Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Baltimore, MD 21224, USA.
authors.forename: Alan B
authors.initials: AB
authors.lastname: Zonderman
authors.affiliation: Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA 02114, USA; Program in Medical and Population Genetics, Broad Institute, Cambridge, MA 02142, USA; Cardiovascular Research Center, Massachusetts General Hospital, Boston, MA 02114, USA; Department of Medicine, Harvard Medical School, Boston, MA 02115, USA.
authors.forename: Sekar
authors.initials: S
authors.lastname: Kathiresan
authors.affiliation: The Novo Nordisk Foundation, Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen 2100, Denmark.
authors.forename: Niels
authors.initials: N
authors.lastname: Grarup
authors.affiliation: Program in Medical and Population Genetics, Broad Institute, Cambridge, MA 02142, USA; Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.
authors.forename: Tõnu
authors.initials: T
authors.lastname: Esko
authors.affiliation: The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA; The Genetics of Obesity and Related Metabolic Traits Program, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA; The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, NY 10069, USA.
authors.forename: Ruth J F
authors.initials: RJ
authors.lastname: Loos
authors.affiliation: Department of Genetics, University of North Carolina, Chapel Hill, NC 27514, USA.
authors.forename: Leslie A
authors.initials: LA
authors.lastname: Lange
authors.affiliation: Department of Anesthesiology & Critical Care Medicine, Johns Hopkins University School of Medicine, Baltimore, MD 21205, USA.
authors.forename: Nauder
authors.initials: N
authors.lastname: Faraday
authors.affiliation: Department of Medicine, Center of Human Nutrition, Washington University School of Medicine, St Louis, MO 63110, USA.
authors.forename: Nada A
authors.initials: NA
authors.lastname: Abumrad
authors.affiliation: Division of Epidemiology, Department of Medicine, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, TN 37235, USA.
authors.forename: Todd L
authors.initials: TL
authors.lastname: Edwards
authors.affiliation: Departments of Internal Medicine and Human Genetics, University of Michigan, Ann Arbor, MI 48108, USA.
authors.forename: Santhi K
authors.initials: SK
authors.lastname: Ganesh
authors.affiliation: Zilber School of Public Health, University of Wisconsin-Milwaukee, Milwaukee, WI 53205, USA.
authors.forename: Paul L
authors.initials: PL
authors.lastname: Auer
authors.affiliation: Population Sciences Branch, National Heart, Lung, and Blood Institute, The Framingham Heart Study, Framingham, MA 01702, USA.
authors.forename: Andrew D
authors.initials: AD
authors.lastname: Johnson
authors.affiliation: Department of Epidemiology, University of Washington, Seattle, WA 98195, USA; Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, WA 98109, USA. Electronic address: apreiner@u.washington.edu.
authors.forename: Alexander P
authors.initials: AP
authors.lastname: Reiner
authors.affiliation: Department of Medicine, Université de Montréal, Montréal, QC H3T 1J4, Canada; Montreal Heart Institute, Montréal, QC H1T 1C8, Canada. Electronic address: guillaume.lettre@umontreal.ca.
authors.forename: Guillaume
authors.initials: G
authors.lastname: Lettre
mesh_terms: D001741:Black or African American / Q000235:genetics; D022981:Allelic Imbalance; D004909:Erythrocyte Indices; D004912:Erythrocytes / Q000166:cytology* / Q000378:metabolism; D004920:Erythropoiesis / Q000235:genetics*; D059472:Exome / Q000235:genetics*; D005787:Gene Frequency; D058685:Genetic Pleiotropy; D014644:Genetic Variation / Q000235:genetics*; D005838:Genotype; D006400:Hematocrit; D006454:Hemoglobins / Q000235:genetics; D006801:Humans; D040641:Quantitative Trait Loci / Q000235:genetics
chemical_list: D006454:Hemoglobins