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SEVENTH EDITION Biochemistry Jeremy M. Berg John L. Tymoczko Lubert Stryer with Gregory J. Gatto, Jr. W. H. Freeman and Company ? New York Publisher: Kate Ahr Parker Developmental Editor: Lisa Samols Senior Project Editor: Georgia Lee Hadler Manuscript Editors: Patricia Zimmerman and Nancy Brooks Design Manager: Vicki Tomaselli Page Make Up: Patrice Sheridan Illustrations: Jeremy Berg with Network Graphics Illustration Coordinator: Janice Donnola Photo Editor: Christine Buese Photo Researcher: Jacalyn Wong Production Coordinator: Paul Rohloff Media Editors: Andrea Gawrylewski, Patrick Shriner, Rohit Phillip, and Marnie Rolfes Supplements Editor: Amanda Dunning Associate Director of Marketing: Debbie Clare Composition: Aptara®, Inc. Printing and Binding: RR Donnelley Library of Congress Control Number: 2010937856 Gregory J. Gatto, Jr., is an employee of GlaxoSmithKline (GSK), which has not supported or funded this work in any way. Any views expressed herein do not necessarily represent the views of GSK. ISBN 13: 9781429229364 ISBN 10: 1429229365 ©2012, 2007, 2002 by W. H. Freeman and Company; © 1995, 1988, 1981, 1975 by Lubert Stryer All rights reserved Printed in the United States of America First printing W. H. Freeman and Company 41 Madison Avenue New York, NY 10010 To our teachers and our students ABOUT THE AUTHORS JEREMY M. BERG received his B.S. and M.S. Tymoczko received his B.A. from the University of degrees in Chemistry from Stanford (where he did Chicago in 1970 and his Ph.D. in Biochemistry from research with Keith Hodgson and Lubert Stryer) and the University of Chicago with Shutsung Liao at the his Ph.D. in Chemistry from Harvard with Richard Ben May Institute for Cancer Research. He then had Holm. He then completed a postdoctoral fellowship a postdoctoral position with Hewson Swift