BIOCHEMISTRY
4TH EDITION
BIOCHEMISTRY DONALD VOET University of Pennsylvania
JUDITH G. VOET Swarthmore College
JOHN WILEY & SONS, INC.
VP & Publisher Kaye Pace Associate Publisher & Editor Petra Recta Sponsoring Editor Joan Kalkut Editorial Assistant Yelena Zolotorevskaya/Patrick White Marketing Manager Kristine Ruff Production Manager Dorothy Sinclair Production Editor Sandra Dumas Senior Designer Madelyn Lesure Senior Illustration Editor Anna Melhorn Executive Media Editor Thomas Kulesa Media Editor Marc Wedzecki Photo Department Manager Hilary Newman Photo Researcher Elyse Rieder Production Management Services Ingrao Associates Cover and part opening art: Illustrations, Irving Geis, Images from Irving Geis Collection/Howard Hughes Medical Institute. Rights owned by HHMI. Not to be reproduced without permission. This book was typeset in 10/12 Times Ten Roman at Aptara®, Inc. and printed and bound by Courier/Kendallville. The cover was printed by Courier/Kendallville. Founded in 1807, John Wiley & Sons, Inc. has been a valued source of knowledge and understanding for more than 200 years, helping people around the world meet their needs and fulfill their aspirations. Our company is built on a foundation of principles that include responsibility to the communities we serve and where we live and work. In 2008, we launched a Corporate Citizenship Initiative, a global effort to address the environmental, social, economic, and ethical challenges we face in our business. Among the issues we are addressing are carbon impact, paper specifications and procurement, ethical conduct within our business and among our vendors, and community and charitable support. For more information, please visit our website: www.wiley.com/go/citizenship. The paper in this book was manufactured by a mill whose forest management programs include sustained yield -harvesting of its timberlands. Sustained yield harvesting principles ensure that the number of trees cut each year does not exceed the amount of new growth. This book is printed on acid-free paper.
Copyright © 2011, 2004, 1995, 1990 by Donald Voet, Judith G Voet. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying recording, scanning or otherwise, except as permitted under Sections 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of the Publisher or authorization through payment of the appropriate per-copy fee to the Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923, (978) 750-8400, fax (978) 646-8600. Requests to the Publisher for permission should be addressed to the Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030-5774, (201) 748-6011, fax (201) 748-6008. Evaluation copies are provided to qualified academics and professionals for review purposes only, for use in their courses during the next academic year. These copies are licensed and may not be sold or transferred to a third party. Upon completion of the review period, please return the evaluation copy to Wiley. Return instructions and a free of charge return shipping label are available at www.wiley.com/go/returnlabel. Outside of the United States, please contact your local representative. ISBN 13 978-0470-57095-1 ISBN 13 978-0470-91745-9 Printed in the United States of America. 10 9 8 7 6 5 4 3 2 1
For our grandchildren: Maya, Leo, Cora, and Elisabeth
ABOUT THE COVER
The cover contains two paintings of horse heart cytochromec. The upper painting, which was drawn by Irving Geis in collaboration with Richard Dickerson, was designed to show the influence of amino acid side chains on the protein’s three-dimensional folding pattern. The lower painting, also made by Geis, is of cytochrome c illuminated by its single iron atom in which its hydrophobic side chains are drawn in green. These paintings were made in the 1970s,
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when only a handful of protein structures were known (around 70,000 are now known) and the personal computers that we presently use to visualize them were many years in the future. It reminds us that biochemistry is a process that is driven by the creativity of the human mind. Our visualization tools have developed from pen, ink, and colored pencils to sophisticated computers and software. Without creativity, however, these tools have little use.
