Biochemical, Physiological, and Molecular Aspects Human Nutrition of This page intentionally left blank THIRD EDITION Biochemical, Physiological, and Molecular Aspects Human Nutrition of Martha H. Stipanuk, PhD Professor Division of Nutritional Sciences Colleges of Human Ecology and Agriculture and Life Sciences Cornell University Ithaca, New York Marie A. Caudill, PhD, RD Associate Professor Division of Nutritional Sciences Colleges of Human Ecology and Agriculture and Life Sciences Cornell University Ithaca, New York 3251 Riverport Lane St. Louis, Missouri 63043 BIOCHEMICAL, PHYSIOLOGICAL, AND MOLECULAR ASPECTS OF HUMAN NUTRITION ISBN: 978-1-4377-0959-9 Copyright © 2013, 2006, 2000 by Saunders, an imprint of Elsevier Inc. All rights reserved. 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To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. Library of Congress Cataloging-in-Publication Data Biochemical, physiological, and molecular aspects of human nutrition/[edited by] Martha H. Stipanuk, Marie A. Caudill. – 3rd ed. p. ; cm. Includes bibliographical references and index. ISBN 978-1-4377-0959-9 (pbk.) I. Stipanuk, Martha H. II. Caudill, Marie A. [DNLM: 1. Nutritional Physiological Phenomena. 2. Metabolism--physiology. QU 145] LC classification not assigned 612.3′9–dc23 2011031519 Senior Editor: Yvonne Alexopoulos Senior Developmental Editors: Lisa P. Newton and Karen C. Turner Editorial Assistant: Kit Blanke Publishing Services Manager: Jeff Patterson Project Manager: Megan Isenberg Designer: Jessica Williams Front cover image copyright Dennis Kunkel Microscopy, Inc. Printed in the United States of America Last digit is the print number: 9 8 7 6 5 4 3 2 1 Chapter Contributors TRACY G. ANTHONY, PhD EDITH BROT-LAROCHE, PhD Associate Professor Research Scientist Department of Biochemistry and Molecular Biology Cordeliers Research Center School of Medicine Department of Physiology, Metabolism, Differentiation Indiana University INSERM/Université Pierre et Marie Curie/CNRS Evansville, Indiana Paris, France RONAL D O. BALL, PhD CHUCK T. CHEN, BSc Professor Emeritus PhD Candidate Department of Agricultural, Food and Nutritional Science Department of Nutritional Sciences University of Alberta University of Toronto Edmonton, Alberta, Canada Toronto, Ontario, Canada RICHARD P. BAZINET, PhD STEVEN K. CLINTON, MD, PhD Assistant Professor Professor Department of Nutritional Sciences Division of Medical Oncology Faculty of Medicine Department of Internal Medicine University of Toronto The Ohio State University Toronto, Ontario, Canada Columbus, Ohio DARLENE E. BERRYMAN, PhD, RD GERALD F. COMBS, Jr., PhD Associate Professor, Nutrition Professor Emeritus School of Applied Health Sciences and Wellness Cornell University Edison Biotechnology Institute Ithaca, New York Ohio University Center Director Athens, Ohio Grand Forks Human Nutrition Research Center USDA-ARS PATSY M. BRANNON, PhD, RD Grand Forks, North Dakota Professor Division of Nutritional Sciences ROBERT R. CRICHTON, PhD, FRSC Cornell University Professor Emeritus Ithaca, New York IMCN, Bâtiment Lavoisier Université Catholique de Louvain J. THOMAS BRENNA, PhD Louvain-la-Neuve, Belgium Professor Division of Nutritional Sciences BRENDA M. DAVY, PhD, RD Cornell University Associate Professor Ithaca, New York Department of Human Nutrition, Foods and Exercise College of Agriculture and Life Sciences JOHN T. BROSNAN, DPhil, DSc Virginia Polytechnic Institute and State University University Research Professor Blacksburg, Virginia Department of Biochemistry Memorial University of Newfoundland HEDLEY C. FREAKE, PhD St. John’s, Newfoundland, Canada Professor Department of Nutritional Sciences MARGARET E. BROSNAN, PhD College of Agriculture and Natural Resources Professor University of Connecticut Department of Biochemistry Storrs, Connecticut Memorial University of Newfoundland St John’s, Newfoundland, Canada v vi Chapter Contributors BALZ FREI, PhD CRYSTAL L. LEVESQUE, PhD Director and Endowed Chair Postdoctoral Fellow Distinguished Professor of Biochemistry and Biophysics Department of Animal & Poultry Science Linus Pauling Institute University of Guelph Oregon State University Guelph, Ontario, Canada Corvallis, Oregon EDWARD O. LIST, PhD JESSE F. GREGORY III, PhD Senior Scientist Professor Edison Biotechnology Institute Department of Food Science and Human Nutrition Ohio University University of Florida Athens, Ohio Gainesville, Florida NELL I. MATTHEWS, BA ARTHUR GRIDER, PhD Research Associate Associate Professor Department of Biochemistry Department of Foods and Nutrition College of Medicine College of Family and Consumer Sciences University of Arkansas for Medical Sciences University of Georgia Little Rock, Arkansas Athens, Georgia MARY M. McGRANE, PhD MATTHEW W. HULVER, PhD Professor—Retired Associate Professor Departments of Nutritional Sciences and Molecular and Department of Human Nutrition, Foods and Exercise Cell Biology College of Agriculture and Life Sciences The University of Connecticut Virginia Polytechnic Institute and State University Storrs, Connecticut Blacksburg, Virginia MARGARET McNURLAN, PhD RONALD J. JANDACEK, PhD Professor of Surgery Adjunct Professor Stony Brook University Medical Center Department of Pathology and Laboratory Medicine Stony Brook, New York University of Cincinnati Cincinnati, Ohio ALEXANDER MICHELS, PhD Research Associate ELIZABETH H. JEFFERY, PhD Linus Pauling Institute Professor Oregon State University Department of Food Science and Human Nutrition Corvallis, Oregon College of Agricultural, Consumer and Environmental Sciences JOSHUA W. MILLER, PhD Department of Pharmacology Professor College of Medicine Department of Medical Pathology and Laboratory Medicine University of Illinois School of Medicine Urbana, Illinois University of California, Davis Sacramento, California ANNA-SIGRID KECK, PhD, CIP Executive Director of the Research Institute DONALD M. MOCK, MD, PhD Carle Foundation Hospital Professor Adjunct Assistant Professor Department of Biochemistry and Molecular Biology Department of Food Science and Human Nutrition Department of Pediatrics University of Illinois College of Medicine Urbana, Illinois University of Arkansas for Medical Sciences Little Rock, Arkansas ARMELLE LETURQUE, PhD Research Scientist PAUL J. MOUGHAN, PhD, DSc, FRSC, FRSNZ Cordeliers Research Center Distinguished Professor and Director Department of Physiology, Metabolism, Differentiation Riddet Institute INSERM/Université Pierre et Marie Curie/CNRS Massey University Paris, France Palmerston North, New Zealand Chapter Contributors vii FORREST H. NIELSEN, PhD HWAI-PING SHENG, PhD Research Nutritionist Honorary Associate Professor U.S. Department Agriculture, Agricultural Research Service Departments of Physiology, Pharmacology, and Pharmacy Grand Forks Human Nutrition Research Center Li Ka Shing Faculty of Medicine Grand Forks, North Dakota The University of Hong Kong Hong Kong, China NOA NOY, PhD Professor JOANNE L. SLAVIN, PhD, RD Departments of Pharmacology and Nutrition Professor Case Western Reserve University Department of Food Science and Nutrition School of Medicine College of Food, Agricultural and Natural Resource Cleveland, Ohio Sciences University of Minnesota SARAH K. ORR, BSc Minneapolis, Minnesota PhD Candidate Department of Nutritional Sciences ARTHUR A. SPECTOR, MD University of Toronto Professor Emeritus Toronto, Ontario, Canada Department of Biochemistry Carver College of Medicine ROBERT S. PARKER, PhD University of Iowa Associate Professor Iowa City, Iowa Division of Nutritional Sciences Cornell University CHRISTINA STARK, MS, RD, CDN Ithaca, New York Nutrition Specialist Division of Nutritional Sciences ELIZABETH N. PEARCE, MD, MSc Cornell University Associate Professor of Medicine Ithaca, New York Section of Endocrinology, Diabetes, and Nutrition Boston University School of Medicine BRUCE R. STEVENS, PhD Boston, Massachusetts Professor Department of Physiology & Functional Genomics W. TODD PENBERTHY, PhD College of Medicine Professor University of Florida Department of Molecular Biology and Microbiology Gainesville, Florida College of Medicine University of Central Florida JUDITH STORCH, PhD Orlando, Florida Professor Department of Nutritional Sciences GAVIN L. SACKS, PhD School of Environmental and Biological Sciences Assistant Professor Rutgers University Department of Food Science New Brunswick, New Jersey Cornell University, New York State Agricultural Experiment Station HEI SOOK SUL, PhD Geneva, New York Professor, Calloway Chair in Human Nutrition Department of Nutritional Science and Toxicology BARRY SHANE, PhD University of California Professor Berkeley, California Department of Nutritional Science and Toxicology University of California KELLY A. TAPPENDEN, PhD, RD Berkeley, California Professor Department of Food Science and Human Nutrition SUE A. SHAPSES, PhD University of Illinois at Urbana-Champaign Professor Champaign, Illinois Department of Nutritional Sciences Rutgers University New Brunswick, New Jersey viii Chapter Contributors CHRISTOPHER A. TAYLOR, PhD, RD, LD ELANGO KATHIRVEL, PhD Associate Professor Research Scientist Medical Dietetics Division and Southern California Institute for Research and Education Department of Family Medicine Long Beach, California School of Allied Medical Professions College of Medicine JOEL B. MASON, MD The Ohio State University Professor of Medicine and Nutrition, Tufts University Columbus, Ohio Scientist I and Director, Vitamins & Carcinogenesis Laboratory, Jean Mayer USDA Human Nutrition ALAN B. R. THOMSON, MD, PhD Research Center on Aging at Tufts University Distinguished University Professor Boston, Massachusetts University of Alberta Adjunct Professor KENGATHEVY MORGAN, PhD University of Western Ontario Research Scientist Canada Southern California Institute for Research and Education Long Beach, California PATRICK TSO, PhD Professor TIMOTHY R. MORGAN, MD Department of Pathology and Laboratory Medicine Chief of Hepatology College of Medicine VA Long Beach Healthcare System University of Cincinnati Long Beach, California Cincinnati, Ohio Professor of Medicine University of California, Irvine JÜRGEN VORMANN, Dr. rer. nat. Irvine, California Director and Professor Institute for Prevention and Nutrition MARGARET P. RAYMAN, DPhil Ismaning/Munich, Germany Professor Division of Nutritional Sciences REIDAR WALLIN, PhD Faculty of Health and Medical Sciences Professor Emeritus University of Surrey Department of Internal Medicine Guildford, United Kingdom Section on Molecular Medicine Wake Forest University School of Medicine KELSEY SHIELDS, BS Winston-Salem, North Carolina Nutritional Sciences Major Division of Nutritional Sciences GARY M. WHITFORD, PhD, DMD Cornell University Regents’ Professor Ithaca, New York Department of Oral Biology College of Dental Medicine PATRICK J. STOVER, PhD Georgia Health Sciences University Professor and Director Augusta, Georgia Division of Nutritional Sciences Cornell University Ithaca, New York CONTRIBUTORS OF NUTRITION INSIGHTS AND CLINICAL CORRELATIONS BARBARA J. STRUPP, PhD Professor SABRINA BARDOWELL, RD Division of Nutritional Sciences and Department PhD Candidate of Psychology Division of Nutritional Sciences Cornell University Cornell University Ithaca, New York Ithaca, New York PATSY M. BRANNON, PhD, RD Professor Division of Nutritional Sciences Cornell University Ithaca, New York Reviewers JOSHUA A. BOMSER, PhD ALISON B. KOHAN, PhD Associate Professor Postdoctoral Fellow Department of Human Nutrition College of Medicine The Ohio State University University of Cincinnati Columbus, Ohio Cincinnati, Ohio SARAH L. BOOTH, PhD STEFANIA LAMON-FAVA, MD, PhD Professor, Friedman School of Nutrition Science and Policy, Associate Professor Tufts University Friedman School of Nutrition Science Senior Scientist and Associate Director, Jean Mayer USDA Tufts University Human Nutrition Research Center on Aging at Tufts Boston, Massachusetts University Boston, Massachusetts KIMBERLY O’BRIEN, PhD Professor PHYLLIS E. BOWEN, PhD, RD Division of Nutritional Sciences Professor Cornell University Department of Kinesiology and Nutrition Ithaca, New York University of Illinois at Chicago Chicago, Illinois MICHAEL E. SPURLOCK, PhD Professor KIMBERLY K. BUHMAN, PhD Department of Food Science and Human Nutrition Assistant Professor Iowa State University Department of Foods and Nutrition Ames, Iowa Purdue University West Lafayette, Indiana JANE ZIEGLER, DCN, RD, LDN, CNSD Department of Nutritional Sciences JOHN E. DOMINY, PhD School of Health Related Professions Postdoctoral Fellow University of Medicine and Dentistry of New Jersey Dana-Farber Cancer Institute Newark, New Jersey Harvard Medical School Boston, Massachusetts CARRIE EARTHMAN, PhD, RD Assistant Professor Department of Food Science and Nutrition University of Minnesota St. Paul, Minnesota ix