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 OXFORD MONOGRAPHS ON MEDICAL GENETICS

General Editors
Arno G. Motulsky Martin Bobrow Peter S. Harper Charles Scriver

Former Editors
J. A. Fraser Roberts C. O. Carter

16. C. R. Scriver and B. Childs: Garrod's inborn factors in disease

18. M. Baraitser: The genetics of neurological disorders
19. R. J. Gorlin, M. M. Cohen, Jr., and L. S. Levin: Syndromes of the head and neck, third edition
21. D. Warburton, J. Byrne, and N. Canki: Chromosome anomalies and prenatal development: an atlas
22. J. J. Nora, K. Berg, and A. H. Nora: Cardiovascular diseases: genetics, epidemiology, and prevention
24. A. E. H. Emery: Duchenne muscular dystrophy, second edition
25. E. G. D. Tuddenham and D. N. Cooper: The molecular genetics of haemostasis and its inherited disorders
26. A. Boue: Fetal medicine
27. R. E. Stevenson, J. G. Hall, and R. M. Goodman: Human malformations
28. R. J. Gorlin, H. V. Toriello, and M. M. Cohen, Jr.: Hereditary hearing loss and its syndromes
29. R. J. M. Gardner and G. R. Sutherland: Chromosome abnormalities and genetic counseling, second edition
30. A. S. Teebi and T. I. Farag: Genetic disorders among Arab populations
31. M. M. Cohen, Jr.: The child with multiple birth defects
32. W. W. Weber: Pharmacogenetics
33. V. P. Sybert: Genetic skin disorders
34. M. Baraitser: Genetics of neurological disorders, third edition
35. H. Ostrer: Non-mendelian genetics in humans
36. E. Traboulsi: Genetic diseases of the eye
37. G. L. Semenza: Transcription factors and human disease
38. L. Pinsky, R. P. Erickson, and R. N. Schimke: Genetic disorders of human sexual development
39. R. E. Stevenson, C. E. Schwartz, and R. J. Schroer: X-linked mental retardation
40. M. J. Khoury, W. Burke, and E. Thomson: Genetics and public health in the 21st century
41. J. Weil: Psychosocial genetic counseling
42. R. J. Gorlin, M. M. Cohen, Jr., and R. C. M. Hennekam: Syndromes of the head and neck, fourth edition
43. M. M. Cohen, Jr., G. Neri, and R. Weksberg: Overgrowth syndromes
44. R. A. King, J. I. Rotter, and A. G. Motulsky: The genetic basis of common diseases, second edition
OXFORD MONOGRAPHS ON MEDICAL GENETICS NO. 44

The Genetic Basis of

Common Diseases
Second Edition

Edited by

RICHARD A. KING
University of Minnesota

JEROME I. ROTTER
Cedars-Sinai Medical Center and UCLA

ARNO G. MOTULSKY
University of Washington

OXPORD
UNIVERSITY PRESS

2002
 OXFORD
UNIVERSITY PRESS

Oxford New York

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and an associated company in Berlin

Copyright © 2002 by Oxford University Press, Inc.

Published by Oxford University Press, Inc.,
198 Madison Avenue, New York, New York, 10016
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Oxford is a registered trademark of Oxford University Press
All rights reserved. No part of this publication may be reproduced,
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electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of Oxford University Press.

Library of Congress Cataloging-in-Publication Data

The genetic basis of common diseases / edited by Richard A. King, Jerome I. Rotter,
Arno G. Motulsky.—2nd ed.
p. ; cm.—(Oxford monographs on medical genetics ; no. 44)
Includes bibliographical references and index.
ISBN 0-19-512582-7
1. Medical genetics. 2. Genetic disorders. I. King, Richard A. (Richard Allen), 1939-
II. Rotter, Jerome I. III. Motulsky, Arno G., 1923- IV. Series.
[DNLM: I. Genetics, Medical. 2. Genetic Diseases, Inborn—genetics. 3. Genetic
Predisposition to Disease. QZ 50 G3226 2002]
RBI55 .G3593 2002
616'.042—dc21
2002018390

1 23456789
Printed in the United States of America
on acid-free paper
 PREFACE TO THE SECOND EDITION

Ten years have elapsed since the first edition of this book ap- To accomplish this goal, the second edition of The Genetic
peared. At that time, the promise of better genetic understand- Basis of Common Diseases has grown from 46 to 55 chapters.
ing of common complex diseases had only recently emerged. More importantly, it has been almost completely rewritten. There
The 1992 volume was a first attempt to describe genetic aspects are 20 entirely new chapters, reflecting advances in diseases
of many of these diseases under one cover. The search for genes whose genetic knowledge was rudimentary at the time of the first
underlying common diseases has now become one of the most edition. In a few cases, chapters have been combined, while other
active areas of biomedical research and is supported by the gov- chapters have been divided. New chapters for specific diseases
ernment and nonprofit foundations as well as by the biotech- cover gallstones, hemochromatosis, osteoporosis, spondy-
nology and pharmaceutical industries. loarthrophies, lupus erythematosis, mental retardation, common
The Human Genome Project has provided an almost com- skin diseases and skin cancer, prostate cancer, hearing loss, and
plete sequence of the human genome and, together with novel migraine. New general chapters address topics such as genetic
methods of population genetics and statistics, is ready to be used counseling, evolution and disease, genetic effects of therapy,
for finding genes involved in complex multigenic diseases. New pharmacogenetics, and the role of mitochondrial variation. The
techniques involving the use of DNA markers such as single nu- authorship is virtually completely changed; close to 90% of the
cleotide polymorphisms (SNPs) are being applied more widely authors (103 out of 117) are new.
and include haplotypic combinations of SNPs in a given chro- This second edition should be useful to readers with a clin-
mosomal region. Knowledge of the genetic basis of common dis- ical, genetic, or epidemiologic orientation who want to learn
ease is expanding and is increasingly brought to the clinic. Cur- more about the genetics of common diseases or who can put
rent excitement must be tempered, however, by the complexity this information to practical use in an increasing number of con-
of the problems: the difficulty in identifying susceptibility genes ditions. We hope that the presentation of current knowledge will
of modest effect, the large scale of the required family studies, lead to novel biomedical and genetic research that brings bet-
the difficulty in proving that a provisionally identified gene is ter understanding, prevention, and treatment of these diseases
indeed responsible for susceptibility, and delineating the mech- that constitute the majority of adult illnesses in the developed
anisms by which implicated genes cause clinical disease. Thus, world.
genetic elucidation of complex diseases has been slower than
some observers had hoped for. While we are learning more about
the role of various genes in complex diseases, the isolation of
specific genes that were localized by linkage studies in complex Minneapolis, Minnesota R.A.K.
diseases has been arduous and remains difficult. It is the goal of Los Angeles, California J.I.R.
this second edition to convey current knowledge, excitement, and Seattle, Washington A.G.M.
balance. June 2002

V
This page intentionally left blank
 CONTENTS

Contributors, ix 12. IgA Deficiency and Common Variable

Immunodeficiency, 210
Harry W. Schroeder, Jr.
Part I Approaches
1. Approach to Genetic Basis of Common Diseases, 3 Part IV Gastrointestinal Disorders
Richard A. King, Jerome I. Rotter, and Arno G. Motulsky
13. Peptic Ulcer and Gastritis, 229
2. Molecular Genetics of Common Disease, 18 A.S. Pena
Jeffrey R. Smith, Thomas D. Gelehrter, and
Francis S. Collins 14. Lactase Deficiency: Biological and Medical Aspects of
the Adult Human Lactase Polymorphism, 250
3. Genetic Epidemiologic Methods, 39 Edward J. Hollox and Dallas M. Swallow
Stephen S. Rich and Thomas A. Sellers
15. Inflammatory Bowel Disease, 266
4. Evolution of Human Genetic Diseases, 50 Huiying Yang, Kent D. Taylor, and Jerome I. Rotter
Jared Diamond and Jerome L Rotter
16. Gallstones, 298
5. Animal Models of Complex Genetic Disease, 65 Beverly Paigen and Martin C. Carey
Aldons J. Lusis, David West, and Richard C. Davis
17. Chronic Liver Disease, 336
6. Genetic Counseling: History, Risk Assessment, Albert J. Czaja
Strategies, and Ethical Considerations, 87
Bonnie S. LeRoy and Ann P. Walker 18. Hereditary Hemochromatosis, 366
Frances Busfield, G.J. Anderson, and L. W. Powell

Part II Cardiopulmonary Diseases 19. Gluten-Sensitive Enteropathy, 382

Susan L. Neuhausen and John J. Zone
7. Genetics of Coronary Atherosclerosis, 105
Arno G. Motulsky and John D. Brunzell Part V Endocrine Disorders
8. Hypertension, 127 20. Thyroid Disease, 397
Steven C. Hunt, Paul N. Hopkins, and Stefan K.G. Grebe
Jean-Marc Lalouel
21. Type 1 Diabetes Mellitus, 431
9. Chronic Obstructive Pulmonary Disease, 155 Leslie J. Raffel and Jerome I. Rotter
Francine Kauffmann and Florence Demenais
22. Type 2 Diabetes Mellitus, 457
Steven C. Elbein, Ken C. Chiu, and M. Alan Permutt
Part III Immunologic and Infectious Diseases
23. Obesity, 481
10. Genetics of Human Susceptibility to Infectious Diseases:
George Bray and Claude Bouchard
Progress and Prospects, 181
Ellen Buschman, Emil Skamene, and Erwin Schurr 24. Genetics of Osteoporosis, 510
Tatiana Foroud, Michael J. Econs,
11. Genetics of Asthma and Bronchial
and C. Conrad Johnston, Jr.
Hyperresponsiveness, 198
Deborah A. Meyers, Denise G. Wiesch, 25. Hyperuricemia and Gout, 518
and Eugene R. Bleecker Michael A. Becker
vii
viii Contents
Part VI Genitourinary Disorders 42. Multiple Sclerosis, 805
Leena Peltonen, Janna Saarela, and Satu Kuokkanen
26. Gynecologic Disorders, 539
Joe Leigh Simpson 43. Alzheimer's Disease, 818
Margaret A. Pericak-Vance and Jonathan L. Haines
27. Infertility and Pregnancy Loss, 554
Carole Ober, M.S. Verp, K.P. Roberts, and J.L. Pry or 44. Affective Disorders, 831
Sevilla D. Detera-Wadleigh and Lynn R. Goldin
Part VII Rheumatologic Diseases 45. Schizophrenia, 850
Ann E. Pulver, Godfrey D. Pearlson, John A. McGrath,
28. Immunology and Immunogenetics, 573 Virginia K. Lasseter, Karen L. Swartz,
Gerald T. Nepom and Henry A. Erlich
and George Papadimitriou
29. Rheumatoid Arthritis, 587 46. Alcoholism, 876
Barbara Nepom and Richard A. King Dharam P. Agarwal
30. Seronegative Spondyloarthropathies, 604 47. Mental Retardation, 914
R. Hal Scofield W. Ted Brown
31. Genetics of Systemic Lupus Erythematosus, 612 48. Hereditary Hearing Loss, 920
Patrick M. Gaffney, Richard A. King,
Rena Ellen Falk and Nathan Fischel-Ghodsian
and Timothy W. Behrens
49. Migraine, 935
32. Genetic Basis of Primary Osteoarthritis, 629 Aarno Palotie, Robert Baloh, and Maija Wessman
John Loughlin and Kay Chapman
33. Common Disorders of Connective Tissue, 638
Reed E. Pyeritz
Part X Other Common Problems
Part VIII Cancer 50. Common Skin Diseases, 951
Virginia P. Sybert
34. Gastrointestinal Cancer, 649
Randall W. Burt 51. Genetic Modulation of Aging and Longevity, 964
George M. Martin
35. Breast Cancer, 670
Katherine L. Nathanson and Barbara L. Weber 52. Mitochondrial Defects in Common Diseases, 975
Douglas C. Wallace and Marie T. Lott
36. Familial and Genetic Influences on Risk of Lung
Cancer, 700 53. Constitutional Chromosome Disorders in Adults, 989
Thomas A. Sellers and Ping Yang Georgia L. Wiesner, David B. Everman,
and Suzanne B. Cassidy
37. Reproductive Organ Cancers, 713
Maren T. Scheuner and Beth Y. Karlan
38. Skin Cancer, 724
Allen E. Bale, Suzanne J. Brown, and William D. Posten Part XI Therapy
39. Prostate Cancer, 738 54. Genetic Consequences of Modern Therapeutics:
William B. Isaacs and Jianfeng Xu latrogenic Mutagenesis, 1023
John J. Mulvihill, Sarah F. Whitton,
40. Hematologic Cancer, 749
and Patrick H. Horn
Marshall Horwitz
55. Pharmacogenetics, Ecogenetics, and
Pharmacogenomics, 1033
Part IX Neuropsychiatric Disorders
Werner Kalow
41. Epilepsy, 779
Magali Fernandez and Thomas D. Bird Index, 1041
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 CONTRIBUTORS

Dharam P. Agarwal, Ph.D., D.Sc. Eugene R. Bleecker, M.D.

Institute of Human Genetics Center for Human Genomics
School of Medicine Department of Medicine
University of Hamburg Wake Forest University School of Medicine
Hamburg, Germany Winston-Salem, North Carolina

Gregory J. Anderson, Ph.D. Claude Bouchard, Ph.D.

Iron Metabolism Laboratory George A. Bray Professor
Queensland Institute of Medical Research Executive Director
Brisbane, Australia Pennington Biomedical Research Center
Louisiana State University
Allen E. Bale, M.D. Baton Rouge, Louisiana
Department of Genetics
Yale University George Bray, M.D.
School of Medicine Boyd Professor
New Haven, Connecticut Pennington Biomedical Research Center
Louisiana State University
Robert Baloh, M.D. Baton Rouge, Louisiana
Department of Neurology
University of California, Los Angeles Suzanne J. Brown, M.D.
School of Medicine Department of Internal Medicine
Los Angeles, California Yale University
School of Medicine
Michael A. Becker, M.D. New Haven, Connecticut
Rheumatology Section
Department of Medicine W. Ted Brown, M.D., Ph.D.
The University of Chicago State University of New York
Pritzker School of Medicine Downstate Medical Center
Chicago, Illinois Brooklyn, New York

Timothy W. Behrens, M.D. John D. Brunzell, M.D.

Department of Medicine Department of Medicine
University of Minnesota Medical School Division of Metabolism, Endocrinology, and Nutrition
Minneapolis, Minnesota University of Washington
School of Medicine
Thomas D. Bird, M.D. Seattle, Washington
Departments of Neurology and Medicine
University of Washington Randall W. Burt, M.D.
School of Medicine Huntsman Cancer Institute
VA Medical Center University of Utah
Seattle, Washington Salt Lake City, Utah

ix
X Contributors
Ellen Buschman, Ph.D. Sevilla D. Detera-Wadleigh, Ph.D.
Department of Medicine Genetics of Mood and Anxiety Unit
McGill University Health Centre Mood and Anxiety Disorders Program
Montreal General Hospital National Institute of Mental Health
Montreal, Quebec National Institutes of Health
Canada Bethesda, Maryland

Frances Busfield, Ph.D. Jared M. Diamond, Ph.D.

Department of Diabetes and Endocrinology Department of Physiology
Princess Alexandra Hospital UCLA School of Medicine
Brisbane, Australia Los Angeles, California

Michael J. Econs, M.D., F.A.C.P., F.A.C.E.

Martin C. Carey, M.D., D.Sc.
Department of Gastroenterology Departments of Medicine and Medical and Molecular Genetics
Indiana University School of Medicine
Brigham and Women's Hospital
Harvard Medical School Indianapolis, Indiana
Boston, Massachusetts Steven C. Elbein, M.D.
Division of Endocrinology
Suzanne B. Cassidy, M.D. University of Arkansas for Medical Sciences
Department of Pediatrics John L. McClellan Veterans Hospital
University of California, Irvine Little Rock, Arkansas
School of Medicine
Irvine, California Henry A. Erlich, Ph.D.
Human Genetics Department
Kay Chapman, Ph.D. Roche Molecular Systems
Institute of Molecular Medicine Alameda, California
University of Oxford
Oxford, United Kingdom David B. Everman, M.D.
Greenwood Genetic Center
Ken C. Chiu, M.D. Greenwood, South Carolina
Division of Endocrinology, Diabetes, and Hypertension
University of California, Los Angeles Rena Ellen Falk, M.D.
School of Medicine Professor of Pediatrics
Los Angeles, California UCLA School of Medicine
Ahmanson Department of Pediatrics
Francis S. Collins, M.D., Ph.D. Medical Genetics and Birth Defects Center
National Human Genome Research Institute Cedar-Sinai Medical Center
National Institutes of Health Los Angeles, California
Bethesda, Maryland
Magali Fernandez, M.D.
Department of Medicine
Albert J. Czaja, M.D.
Ohio State University
Division of Gastroenterology and Hepatology
College of Medicine
Department of Medicine
Columbus, Ohio
Mayo Clinic and Mayo Foundation
Rochester, Minnesota Nathan Fischel-Ghodsian, M.D.
Professor of Pediatrics
Richard C. Davis, Ph.D. UCLA School of Medicine
Department of Medicine Ahmanson Department of Pediatrics
University of California, Los Angeles Medical Genetics and Birth Defects Center
School of Medicine Cedar-Sinai Medical Center
Los Angeles, California Los Angeles, California

Florence Demenais, M.D. Tatiana Foroud, Ph.D.

Department of Statistical Genetics and Departments of Medical and Molecular Genetics and
Genetic Epidemiology of Multifactorial Diseases Psychiatry
INSERM EMI 00-06 Indiana University School of Medicine
Evry, France Indianapolis, Indiana
 Contributors xi
Patrick M. Gaffney, M.D. William B. Isaacs, Ph.D.
Department of Medicine Department of Urology
University of Minnesota Medical School Johns Hopkins University
Minneapolis, Minnesota School of Medicine
Baltimore, Maryland
Thomas D. Gelehrter, M.D.
Department of Human Genetics C. Conrad Johnston, Jr., M.D.
University of Michigan Department of Medicine
Ann Arbor, Michigan Indiana University School of Medicine
Indianapolis, Indiana
Lynn R. Goldin, Ph.D. Werner Kalow, M.D.
Genetic Epidemiology Branch
Department of Pharmacology
Division of Cancer Epidemiology and Genetics
University of Toronto
National Cancer Institute
Toronto, Ontario
National Institutes of Health
Canada
Bethesda, Maryland
Beth Y. Karlan, M.D.
Stefan K.G. Grebe, Arzt, M.D., F.R.A.C.P. Department of Obstetrics and Gynecology
Department of Pathology and Molecular Medicine University of California, Los Angeles
University of Otago School of Medicine
Wellington School of Medicine and Health Sciences Los Angeles, California
Wellington, New Zealand
Francine Kauffman, M.D.
Jonathan L. Haines, Ph.D. Department of Epidemiology and Biostatistics
Department of Molecular Physiology and Biophysics INSERM U472
Program in Human Genetics Villejuif, France
Vanderbilt University Medical Center
Richard A. King, M.D., Ph.D.
Nashville, Tennessee
Department of Medicine
University of Minnesota Medical School
Edward J. Hollox, Ph.D.
Minneapolis, Minnesota
Institute of Genetics
University of Nottingham Satu Kuokkanen, M.D., Ph.D.
Nottingham, United Kingdom Department of Obstetrics and Gynecology
Columbia University
Paul N. Hopkins, M.D., M.S.P.M. College of Physicians and Surgeons
Department of Internal Medicine New York, New York
University of Utah
School of Medicine Jean-Marc Lalouel, M.D., D.Sc.
Salt Lake City, Utah Department of Human Genetics
University of Utah
Patrick H. Horn, B.S. School of Medicine
Department of Pediatrics Salt Lake City, Utah
University of Oklahoma
Virginia K. Lasseter, B.A.
College of Medicine
Department of Psychiatry
Oklahoma City, Oklahoma
Johns Hopkins University
Marshall Horwitz, M.D., Ph.D. School of Medicine
Division of Medical Genetics Baltimore, Maryland
Department of Medicine Bonnie S. LeRoy, M.S.
University of Washington Department of Genetics, Cell Biology, and Development
School of Medicine The University of Minnesota
Seattle, Washington Minneapolis, Minnesota
Steven C. Hunt, Ph.D. Marie T. Lott, M.A.
Department of Internal Medicine Center for Molecular Medicine
University of Utah Emory University
School of Medicine School of Medicine
Salt Lake City, Utah Atlanta, Georgia
xli Contributors
John Loughlin, Ph.D. Susan L. Neuhausen, Ph.D.
Institute of Molecular Medicine Department of Medical Informatics
University of Oxford University of Utah School of Medicine
Oxford, United Kingdom Salt Lake City, Utah

Aldons J. Lusis, Ph.D. Carole Ober, Ph.D.

Departments of Medicine, of Microbiology, Immunology, Departments of Human Genetics
and Molecular Genetics, and of Human Genetics and Obstetrics and Gynecology
University of California, Los Angeles The University of Chicago
School of Medicine Chicago, Illinois
Los Angeles, California
Beverly Paigen, Ph.D.
George M. Martin, M.D. The Jackson Laboratory
Departments of Pathology and Genome Sciences Bar Harbor, Maine
University of Washington
School of Medicine Aarno Palotie, M.D., Ph.D.
Seattle, Washington Department of Pathology and Laboratory Medicine
University of California, Los Angeles
John A. McGrath, M.S. School of Medicine
Department of Psychiatry Los Angeles, California
Johns Hopkins University
School of Medicine George Papadimitriou, M.D.
Baltimore, Maryland University Mental Health Research Institute
Eginition Hospital
Deborah A. Meyers, Ph.D.
Athens, Greece
Center for Human Genomics
Departments of Pediatrics and Public Health Godfrey D. Pearlson, M.D.
Wake Forest University School of Medicine Department of Psychiatry
Winston-Salem, North Carolina Johns Hopkins University
School of Medicine
Arno G. Motulsky, M.D.
Baltimore, Maryland
Departments of Medicine (Division of Medical Genetics)
and Genome Sciences
Leena Peltonen, M.D., Ph.D.
University of Washington
Department of Human Genetics
School of Medicine
University of California, Los Angeles
Seattle, Washington
School of Medicine
John J. M u l v i h i l l , M.D. Los Angeles, California
Department of Pediatrics Biomedicum, University of Helsinki
University of Oklahoma National Public Health Institute
College of Medicine Helsinki, Finland
Oklahoma City, Oklahoma
A.S. Pena, M.D., Ph.D., F.R.C.P.
Katherine L. Nathanson, M.D. Professor of Gastrointestinal Immunology
Department of Medicine Head Laboratory of Immunogenetics
University of Pennsylvania Department of Gastroenterology
School of Medicine Vrije Universiteit Medical Center
Philadelphia, Pennsylvania Amsterdam, The Netherlands

Barbara Nepom, M.D. Margaret A. Pericak-Vance, Ph.D.

Immunology Program Center for Human Genetics
Virginia Mason Research Center Department of Medicine
Seattle, Washington Duke University Medical Center
Durham, North Carolina
Gerald T. Nepom, M.D., Ph.D.
Virginia Mason Research Center and M. Alan Permutt, M.D.
Department of Immunology Division of Endocrinology, Diabetes, and Metabolism
University of Washington School of Medicine Washington University School of Medicine
Seattle, Washington St. Louis, Missouri
 Contributors xiii
W i l l i a m D. Posten, M.D. Maren T. Scheuner, M.D., M.P.H.
Department of Genetics Department of Medicine
Yale University University of California, Los Angeles
School of Medicine School of Medicine
New Haven, Connecticut Los Angeles, California
Lawrie W. Powell, M.D., Ph.D. Harry W. Schroeder, Jr., M.D., Ph.D., F.A.C.M.G.
Iron Metabolism Laboratory Division of Developmental and Clinical Immunology
Queensland Institute of Medical Research Comprehensive Cancer Center
Brisbane, Australia Departments of Medicine and Microbiology
Jon L. Pryor, M.D., Ph.D. University of Alabama at Birmingham
Department of Urologic Surgery Birmingham, Alabama
University of Minnesota Medical School
Minneapolis, Maryland Erwin Schurr, Ph.D.
Departments of Medicine and Human Genetics
Ann E. Pulver, Sc.D. McGill Centre for the Study of Host Resistance
Department of Psychiatry Montreal General Hospital
Johns Hopkins University Montreal, Quebec
School of Medicine Canada
Baltimore, Maryland
R. Hal Scofield, M.D.
Reed E. Pyeritz, M.D., Ph.D. Department of Medicine
Division of Medical Genetics University of Oklahoma
Department of Medicine College of Medicine
University of Pennsylvania School of Medicine Arthritis and Immunology Program
Philadelphia, Pennsylvania Oklahoma Medical Research Foundation
Leslie J. Raffel, M.D. Oklahoma City, Oklahoma
Medical Genetics—Birth Defects Center
Department of Pediatrics Thomas A. Sellers, Ph.D., M.P.H.
Cedars-Sinai Medical Center Department of Health Sciences Research
Department of Pediatrics Mayo Clinic and Foundation
University of California, Los Angeles Rochester, Minnesota
School of Medicine
Los Angeles, California Joe Leigh Simpson, M.D.
Department of Obstetrics and Gynecology
Stephen S. Rich, Ph.D. Baylor College of Medicine
Department of Public Health Sciences Houston, Texas
Wake Forest University
School of Medicine Emil Skamene, M.D., Ph.D.
Winston-Salem, North Carolina Departments of Medicine and Human Genetics
McGill University Health Centre
Kenneth P. Roberts, Ph.D. Montreal General Hospital
Departments of Urologic Surgery and Genetics, Cell Biology, Montreal, Quebec
and Development Canada
University of Minnesota Medical School
Minneapolis, Minnesota Jeffrey R. Smith, M.D., Ph.D.
Jerome I. Rotter, M.D. Department of Medicine
Director, Division of Medical Genetics Division of Genetic Medicine
Cedars-Sinai Board of Governors' Chair in Medical Genetics Vanderbilt University Medical Center
Departments of Medicine, Pediatrics, and Human Genetics Nashville, Tennessee
University of California, Los Angeles
Los Angeles, California Dallas M. Swallow, Ph.D.
Department of Human Genetics
Janna Saarela, M.D., Ph.D. The Gallon Laboratory
Biomedicum, University of Helsinki Department of Biology
National Public Health Institute University College London
Helsinki, Finland London, United Kingdom
xiv Contributors
Karen L. Swartz, M.D. David West, Ph.D.
Department of Psychiatry Xenogen
Johns Hopkins University Alameda, California
School of Medicine
Baltimore, Maryland Sarah F. Whitton
Department Pediatrics
Virginia P. Sybert, M.D. University of Oklahoma
Division of Dermatology College of Medicine
Department of Medicine Oklahoma City, Oklahoma
University of Washington
School of Medicine Denise G. Wiesch, Ph.D.
Seattle, Washington Division of Allergy, Immunology, and Transplantation
National Institute of Allergy and Infectious Diseases
Kent D. Taylor, Ph.D. National Institutes of Health
Medical Genetics Birth Defects Center Bethesda, Maryland
Imflammatory Bowel Disease Center
Departments of Medicine and Pediatrics Georgia L. Wiesner, M.D.
Steven Spielberg Pediatric Research Center Departments of Genetics and Medicine
Cedars-Sinai Research Institute Case Western Reserve University
University of California, Los Angeles School of Medicine and
School of Medicine University Hospitals of Cleveland
Los Angeles, California Cleveland, Ohio
Marian S. Verp, M.D. J i a n f e n g X u , M.D., M.P.H.
Departments of Obstetrics and Gynecology Department of Public Health
And Human Genetics Center for Human Genomics
The University of Chicago Wake Forest University
Chicago, Illinois School of Medicine
Winston-Salem, North Carolina
Ann P. Walker, M.A.
Department of Pediatrics H u i y i n g Yang, M . D . , Ph.D.
University of California, Irvine Medical Genetics Birth Defects Center
Orange, California Inflammatory Bowel Disease Center
Douglas C. Wallace, Ph.D. Departments of Medicine and Pediatrics
Center for Molecular Medicine Steven Spielberg Pediatric Research Center
Emory University Cedars-Sinai Research Institute
School of Medicine University of California, Los Angeles
Atlanta, Georgia School of Medicine
Los Angeles, California
Barbara L. Weber, M.D.
Department of Medicine and Genetics Ping Yang, M.D., Ph.D.
University of Pennsylvania Department of Health Sciences Research
School of Medicine Mayo Clinic
Philadelphia, Pennsylvania Rochester, Minnesota

Maija Wessman, Ph.D. John J. Zone, M.D.

Department of Biosciences Department of Dermatology
University of Helsinki University of Utah School of Medicine
Helsinki, Finland Salt Lake City, Utah
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