Folate in Health and Disease, 2nd Edition

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 Contents
Preface.......................................................................................................................ix
Editor ........................................................................................................................xi
Contributors ........................................................................................................... xiii

Chapter 1 Folate Chemistry and Metabolism ....................................................... 1

Barry Shane

Chapter 2 Folate Bioavailability .........................................................................25

Helene McNulty and Kristina Pentieva

Chapter 3 Folate Biochemical Pathways and Their Regulation.......................... 49

Patrick J. Stover

Chapter 4 Genetic Variation: Effect on Folate Metabolism

and Health .......................................................................................... 75
Karen E. Christensen and Rima Rozen

Chapter 5 Folate in Pregnancy and Lactation ................................................... 111

Tsunenobu Tamura, Mary Frances Picciano, and
Michelle Kay McGuire

Chapter 6 Folate Status and Birth Defect Risk: Epidemiological

Perspective........................................................................................ 133
Charlotte A. Hobbs, Gary M. Shaw, Martha M. Werler, and
Bridget Mosley

Chapter 7 Folate-Related Birth Defects: Embryonic Consequences of

Abnormal Folate Transport and Metabolism ................................... 155
Deeann Wallis, Johnathan L. Ballard, Gary M. Shaw,
Edward J. Lammer, and Richard H. Finnell

Chapter 8 Folic Acid Fortification: Neural Tube Defect

Risk Reduction—A Global Perspective .......................................... 179
Robert J. Berry, Joseph Mulinare, and Heather C. Hamner

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Chapter 9 Folate and Cancer: Epidemiological Perspective .............................205

Jia Chen, Xinran Xu, Amy Liu, and Cornelia M. Ulrich

Chapter 10 Folate and Carcinogenesis: Basic Mechanisms................................ 235

Eric Ciappio and Joel B. Mason

Chapter 11 Folate and Vascular Disease: Epidemiological Perspective ............. 263

Sari R. Kalin and Eric B. Rimm

Chapter 12 Folate and Vascular Disease: Basic Mechanisms ............................ 291

Luciana Hannibal, Alla V. Glushchenko, and Donald W. Jacobsen

Chapter 13 Folate and Neurological Function: Epidemiological Perspective .... 325

Martha Savaria Morris and Paul F. Jacques

Chapter 14 Folate and Neurological Disease: Basic Mechanisms ...................... 355

Teodoro Bottiglieri and Edward Reynolds

Chapter 15 Folate–Vitamin B12 Interrelationships: Links to Disease Risk ....... 381

Anne M. Molloy

Chapter 16 Clinical Folate Deficiency ................................................................409

Sally P. Stabler

Chapter 17 Influence of Alcohol on Folate Status and Methionine

Metabolism in Relation to Alcoholic Liver Disease ....................... 429
Charles H. Halsted, Valentina Medici, and Farah Esfandiari

Chapter 18 Folate and Choline Interrelationships: Metabolic and

Potential Health Implications ...........................................................449
Marie A. Caudill

Chapter 19 Folate: Recommended Intakes, Consumption, and Status .............. 467

Gail P. A. Kauwell, Megan L. Diaz, Quanhe Yang,
and Lynn B. Bailey

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 Contents vii

Chapter 20 Kinetics of Folate and One-Carbon Metabolism ............................. 491

Jesse F. Gregory III, Vanessa R. da Silva, and Yvonne Lamers

Chapter 21 Folate Analytical Methodology........................................................ 517

Christine M. Pfeiffer, Zia Fazili, and Mindy Zhang

Index ...................................................................................................................... 575

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 Preface
More than a decade has passed since the first edition of Folate in Health and Disease
was published. During this time there have been thousands of new research stud-
ies related to folate and its link to disease and birth defect risk, thus providing the
impetus for an updated interpretation of this large body of scientific evidence. The
public health implications of these new findings are enormous; therefore, the second
edition of Folate in Health and Disease bridges basic science with clinical medicine
and public health.
The first several chapters in the new edition are organized to provide the reader
with background knowledge related to folate chemistry, metabolism, bioavailability,
and the influence of genetic polymorphisms. Folate’s role in reproduction and birth
defect prevention is then reviewed, followed by a separate chapter in which epi-
demiological evidence linking specific birth defects and folate status is evaluated.
Intriguing research findings related to basic mechanisms provide insight regarding
potential etiologies of folate-responsive birth defects. Folic acid fortification, which
was implemented after the first edition of this book was published, is described from
a global perspective as it relates to neural tube defect risk reduction.
Chronic disease is covered in a similar manner to that of birth defects; epidemio-
logical data are first critiqued in separate chapters related to cancer, vascular, and
neurological diseases, coupled with subsequent chapters in which proposed mecha-
nisms are presented. The potential for folic acid to affect mortality and morbidity
associated with the recurrence of vascular disease and cancer has been the focus
of ongoing controlled intervention trials. This new edition of Folate in Health and
Disease is timely in that the findings from these studies can now be described and
interpreted to benefit investigators in both the clinical/public health and basic sci-
ence arenas.
The interrelationships between folate and other nutrients required for normal one-
carbon metabolism are covered in several chapters, and the biochemical and clinical
ramifications of alterations in status are highlighted. The interaction between folate
and vitamin B12 is addressed from a biochemical and public health perspective.
The complexities of diagnosis and treatment of a clinical folate deficiency are dis-
cussed, followed by a related chapter on the effect of alcohol on folate and methi-
onine metabolism. Choline is covered in a separate chapter that presents evidence
related to the metabolic interaction between folate and choline and the implications
for health maintenance.
Dietary intake recommendations for select countries worldwide are compared
with an overview of the approaches used by the Institute of Medicine’s commit-
tee to estimate the Dietary Reference Intakes. Changes in folate status over time
within the US population are a focus of this chapter with attention given to the
influence of folic acid fortification and supplement use on folate status. Estimated
dietary folate intakes for the US population and specific population subgroups are
presented.

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 x Preface

A critique of experimental approaches and an evaluation of studies of one-carbon

metabolism provide a new understanding of the knowledge regarding the rates of
folate utilization and how to best investigate factors that may influence these rates.
Established and new methods for the measurement of folate/folic acid in food and
physiological fluids are critiqued in the final chapter as a means to evaluate and com-
pare results from studies in which different methodologies were used. In addition
to providing insight into the interpretation of published data, this chapter provides
research scientists with guidelines regarding the most appropriate method for future
research studies.
In summary, the second edition of Folate in Health and Disease integrates and
provides an interpretation of basic and applied knowledge related to folate and its role
in health maintenance and disease prevention. The interdisciplinary approach taken
in this new book results in a valuable resource for both basic and applied research
scientists and nutritionists in academics, clinical medicine, and public health.

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 Editor
Lynn B. Bailey, PhD, is a professor in the Food Science and Human Nutrition
Department, University of Florida, Gainesville, Florida. Before joining the faculty
at the University of Florida, she received a PhD from Purdue University, an MS from
Clemson University, and a BS from Winthrop University. The focus of Dr. Bailey’s
research program is the estimation of folate requirements involving metabolic stud-
ies conducted with human subjects of all ages. Evaluation of the impact of genetic
variants on folate requirements and biomarkers for disease and birth defect risk is a
key research emphasis.
Dr. Bailey has served as a member of the Institute of Medicine’s Dietary Reference
Intake committee for folate, vitamin 12, and other B vitamins, and she was a mem-
ber of the Food and Drug Administration’s Folic Acid Advisory Committee. She has
served as a scientific advisor for the Centers for Disease Control and Birth Defect
Prevention, as well as for other organizations, including the March of Dimes and
the Pan American Health Organization, on projects focused on neural tube defect
prevention in the United States and developing countries. Dr. Bailey has received
numerous awards, including the USDA Superior Achievement Award; the March of
Dimes’ Agnes Higgins Award, Maternal-Fetal Nutrition; the American Society for
Nutrition’s Centrum Science Award for influence on current scientific knowledge of
folate status and requirements of humans; and the University of Florida’s Teacher
Scholar of the Year award.

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 Contributors
Lynn B. Bailey, PhD
Professor, Department of Food Science and Human Nutrition, University of Florida,
Gainesville, Florida

Johnathan L. Ballard, PhD

Assistant Research Scientist, Center for Environmental and Genetic Medicine,
Institute of Biosciences and Technology, Texas A&M Health Science Center,
Houston, Texas

Robert J. Berry, MD, MPHTM

Medical Epidemiologist, Birth Defects Epidemiology Team Leader, National Center
on Birth Defects and Developmental Disabilities, Centers for Disease Control and
Prevention, Atlanta, Georgia

Teodoro Bottiglieri, PhD

Director of Neuropharmacology Laboratory, Adjunct Professor of Biomedical
Studies, Baylor University Medical Center, Institute of Metabolic Disease, Dallas,
Texas

Marie A. Caudill, PhD, RD

Associate Professor, Division of Nutritional Sciences, Cornell University, Ithaca,
New York

Jia Chen, ScD

Associate Professor, Departments of Community and Preventive Medicine, Pediatrics
and Oncological Science, Mount Sinai School of Medicine, New York, New York

Karen E. Christensen, PhD

Postdoctoral Fellow, Departments of Human Genetics and Pediatrics, McGill
University, Montreal, Quebec, Canada

Eric Ciappio, MS, RD

Doctoral Student, Friedman School of Nutrition Science Policy, Tufts University;
Vitamins & Carcinogenesis Laboratory, Jean Mayer USDA Human Nutrition
Research Center on Aging at Tufts University, Boston, Massachusetts

Vanessa R. da Silva, BS
Doctoral Student, Department of Food Science and Human Nutrition, University of
Florida, Gainesville, Florida

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 xiv Contributors

Megan L. Diaz, MS, RD, LD

Graduate Assistant, University of Florida, Gainesville, Florida

Farah Esfandiari, PhD

Research Faculty, Department of Internal Medicine, University of California Davis,
Davis, California

Zia Fazili, MPhil, PhD

Research Chemist, Nutritional Biomarkers Branch, Division of Laboratory Sciences,
Centers for Disease Control and Prevention, Atlanta, Georgia

Richard H. Finnell, PhD

Regents Professor, Center for Environmental and Genetic Medicine, Institute
of Biosciences and Technology, Texas A&M Health Science Center, Houston,
Texas

Alla V. Glushchenko, MD, PhD

Postdoctoral Fellow, Department of Cell Biology, Lerner Research Institute,
Cleveland Clinic, Cleveland, Ohio

Jesse F. Gregory III, PhD

Professor, Department of Food Science and Human Nutrition, University of Florida,
Gainesville, Florida

Charles H. Halsted, MD
Professor, Departments of Internal Medicine and Nutrition, University of California
Davis, Davis, California

Heather C. Hamner, MS, MPH

Nutrition Epidemiologist, Division of Birth Defects and Developmental Disabilities,
National Center on Birth Defects and Developmental Disabilities, Centers for Disease
Control and Prevention, Atlanta, Georgia

Luciana Hannibal, BSc

Doctoral Student, Department of Cell Biology, Lerner Research Institute, Cleveland
Clinic, Cleveland, Ohio; and School of Biomedical Sciences, Kent State University,
Kent, Ohio

Charlotte A. Hobbs, MD, PhD

Professor, Department of Pediatrics, College of Medicine, University of Arkansas
for Medical Sciences, Little Rock, Arkansas

Donald W. Jacobsen, PhD, FAHA

Staff, Department of Cell Biology, Lerner Research Institute, and Department of
Cardiovascular Medicine, Heart and Vascular Institute, Cleveland Clinic, Cleveland,
Ohio; Professor of Molecular Medicine, Department of Molecular Medicine,

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 Contributors xv

Cleveland Clinic Lerner College of Medicine, Case Western Reserve University,

Cleveland, Ohio

Paul F. Jacques, ScD

Senior Scientist and Director, Nutritional Epidemiology, Jean Myer USDA Human
Nutrition Research Center on Aging, and Professor, Friedman School of Nutrition
Science and Policy, Tufts University, Boston, Massachusetts

Sari R. Kalin, MS, RD, LDN

Program Coordinator, Department of Nutrition, Harvard School of Public Health,
Boston, Massachusetts

Gail P. A. Kauwell, PhD, RD, LDN

Professor, Food Science and Human Nutrition Department, University of Florida,
Gainesville, Florida

Yvonne Lamers, PhD

Postdoctoral Associate, Department of Food Science and Human Nutrition,
University of Florida, Gainesville, Florida

Edward J. Lammer, MD
Associate Scientist, Children’s Hospital Oakland Research Institute, Oakland,
California

Amy Liu, MPH

Doctoral Student, Department of Epidemiology, University of Washington, Seattle,
Washington

Joel B. Mason, MD
Associate Professor, Schools of Medicine and Nutritional Science & Policy,
Tufts University; Director, Vitamins & Carcinogenesis Laboratory, Jean Mayer
USDA Human Nutrition Research Center on Aging at Tufts University, Boston,
Massachusetts

Michelle Kay McGuire, PhD

Associate Professor, School of Molecular Biosciences, Washington State University,
Pullman, Washington

Helene McNulty, PhD, RD

Professor, Northern Ireland Centre for Food and Health, School of Biomedical
Sciences, University of Ulster, Coleraine, Northern Ireland

Valentina Medici, MD
Assistant Professor, Department of Internal Medicine, Division of Gastroenterology
and Hepatology, University of California Davis Medical Center, Sacramento,
California

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 xvi Contributors

Anne Molloy, PhD

Research Senior Lecturer, Department of Clinical Medicine, School of Medicine,
Trinity College, Dublin, Ireland

Martha Savaria Morris, PhD

Epidemiologist, Nutritional Epidemiology Program, Jean Mayer USDA Human
Nutrition Research Center on Aging at Tufts University, Boston, Massachusetts

Bridget S. Mosley, MPH

Epidemiologist, Department of Pediatrics, College of Medicine, University of
Arkansas for Medical Sciences, Little Rock, Arkansas

Joseph Mulinare, MD, MSPH

Chief, Prevention Research & Health Communications Team, National Center on
Birth Defects and Developmental Disabilities, Centers for Disease Control and
Prevention, Atlanta, Georgia

Kristina Pentieva, MD, PhD

Senior Lecturer, Northern Ireland Centre for Food and Health, School of Biomedical
Sciences, University of Ulster, Coleraine, Northern Ireland

Christine M. Pfeiffer, PhD

Chief of Nutritional Biomarkers Branch, Division of Laboratory Sciences, Centers
for Disease Control and Prevention, Atlanta, Georgia

Mary Frances Picciano, PhD

Senior Nutrition Research Scientist, Office of Dietary Supplements, National
Institutes of Health, Bethesda, Maryland

Edward Reynolds, MD, FRCP, FRCPsych

Honorary Senior Lecturer, Department of Neurology, King’s College, University of
London, London, England

Eric B. Rimm, ScD

Associate Professor of Medicine, Harvard Medical School, Channing Laboratory,
Brigham and Women’s Hospital; Associate Professor of Epidemiology and Nutrition,
Director, Program in Cardiovascular Epidemiology, Harvard School of Public
Health, Boston, Massachusetts

Rima Rozen, PhD, FCCMG

James McGill Professor, Departments of Human Genetics and Pediatrics, McGill
University, Montreal, Quebec, Canada

Barry Shane, PhD

Professor, Department of Nutritional Sciences and Toxicology, University of
California, Berkeley, California

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 Contributors xvii

Gary M. Shaw, DrPH

Professor, Department of Pediatrics, Stanford University, Palo Alto, California

Sally P. Stabler, MD
Co-Division Head of Hematology, Professor of Medicine, Department of Medicine,
University of Colorado at Denver, Aurora, Colorado

Patrick J. Stover, PhD

Professor of Nutritional Biochemistry, Division of Nutritional Sciences, Cornell
University, Ithaca, New York

Tsunenobu Tamura, MD
Professor, Department of Nutrition Sciences, University of Alabama at Birmingham,
Birmingham, Alabama

Cornelia M. Ulrich, PhD

Full Member, Cancer Prevention Program, Fred Hutchinson Cancer Research
Center, Seattle, Washington; Head, Division of Preventive Oncology, German Cancer
Research Center, and Co-Director, National Center for Tumor Diseases, Heidelberg,
Germany

Deeann Wallis, PhD

Research Scientist, Texas A&M Institute for Genomic Medicine, Texas A&M Health
Science Center, Houston, Texas

Martha M. Werler, ScD

Professor, Epidemiology, Slone Epidemiology Center at Boston University, Boston,
Massachusetts

Xinran Xu, PhD

Research Fellow, Department of Community and Preventive Medicine, Mount Sinai
School of Medicine, New York, New York

Quanhe Yang, PhD

Epidemiologist, Office of Public Health Genomics, Centers for Disease Control and
Prevention, Atlanta, Georgia

Ming Zhang, MD
Research Microbiologist, Nutritional Biomarkers Branch, Division of Laboratory
Sciences, Centers for Disease Control and Prevention, Atlanta, Georgia

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