Biochemistry, Molecular Biology, and Genetics Todd A. Swanson, M.D., Ph.D. Resident in Radiation Oncology William Beaumont Hospital Royal Oak, Michigan Sandra I. Kim, M.D., Ph.D. Division of Nuclear Medicine and Molecular Imaging Massachusetts General Hospital Boston, Massachusetts Marc J. Glucksman, Ph.D. Professor, Department of Biochemistry and Molecular Biology Director, Midwest Proteome Center Rosalind Franklin University of Medicine and Science The Chicago Medical School North Chicago, Illinois WITH EDITORIAL CONSULTATION BY Michael A. Lieberman, Ph.D. Dean, Instructional and Research Computing, UCit Distinguished Teaching Professor University of Cincinnati Cincinnati, OH Acquisitions Editor: Charles W. Mitchell Product Manager: Stacey L. Sebring Marketing Manager: Jennifer Kuklinski Designer: Holly Reid McLaughlin Compositor: Cadmus Communications Printer: C & C Offset Printing Copyright C 2010 Lippincott Williams &Wilkins 351 West Camden Street Baltimore, MD 21201 530 Walnut Street Philadelphia, PA 19106 All rights reserved. This book is protected by copyright. No part of this book may be reproduced in any form or by any means, including photocopying, or utilized by any information storage and retrieval system without written permission from the copyright owner. The publisher is not responsible (as a matter of product liability, negligence, or otherwise) for any injury resulting from any material contained herein. This publication contains information relating to general principles of medical care that should not be construed as specific instructions for individual patients. Manufacturers’ product information and package inserts should be reviewed for current information, including contraindications, dosages, and precautions. Printed in Hong Kong First Edition, 1990 Second Edition, 1994 Third Edition, 1999 Fourth Edition, 2007 Library of Congress Cataloging-in-Publication Data Swanson, Todd A. Biochemistry, molecular biology, and genetics / Todd A. Swanson, Sandra I. Kim, Marc J. Glucksman ; with editorial consultation by Michael A. Lieberman. — 5th ed. p. ;cm. — (Board review series) Rev. ed. of: Biochemistry and molecular biology / Todd A. Swanson, Sandra I. Kim, Marc J. Glucksman. 4th ed. c2007. Includes bibliographical references and index. ISBN 978-0-7817-9875-4 (hardcopy : alk. paper) 1. Biochemistry—Examinations, questions, etc. 2. Molecular biology—Examinations, questions, etc. I. Kim, Sandra I. II. Glucksman, Marc J. III. Lieberman, Michael, 1950- IV. Swanson, Todd A. Biochemistry and molecular biology. V. Title. VI. Series: Board review series. [DNLM: 1. Biochemical Phenomena—Examination Questions. 2. Biochemical Phenomena—Outlines. 3. Genetic Processes—Examination Questions. 4. Genetic Processes—Outlines. QU 18.2 S972b 2010] QP518.3.S93 2010 572.8076—dc22 2009029693 The publishers have made every effort to trace the copyright holders for borrowed material. If they have inadvertently overlooked any, they will be pleased to make the necessary arrangements at the first opportunity. To purchase additional copies of this book, call our customer service department at (800) 638-3030 or fax orders to (301) 223-2320. International customers should call (301) 223-2300. Visit Lippincott Williams &Wilkins on the Internet: http://www.LWW.com. Lippincott Williams &Wilkins customer service representatives are available from 8:30 am to 6:00 pm, EST. For Olga, Maxwell, Anneliese, and the eagerly awaited new addition to the Swanson clan. If not for you, all my efforts would be in vain. Preface This revision of BRS Biochemistry, Molecular Biology, and Genetics includes additional high-yield material to help the reader master clinical principles of medical biochemistry as they prepare for the revamped Step 1 USMLE. Our goal is to offer a review book that both lays the foundations of biochemistry and introduces clinically relevant correlates. In doing so, we have de-emphasized some of the rote memorization of structures and formulas that often obscure the big picture of medical biochemistry. Clinical Correlates in each chapter provide additional clinical insight, distilling numerous clinical correlations into a format that offers the highest yield in review. We hope that these correlations will help answer a commonly asked question: ‘‘Why do we have to know this for the boards?’’ This revised edition also includes a new chapter on genetics as related to medical bio- chemistry. We hope this chapter will augment other review texts on genetics that students may consult in preparation for Step 1. Many of the questions at the end of each chapter have been revised to maximize their value for the student preparing for the exam. A comprehensive exam at the end of this vol- ume reinforces the concepts of the text. Our objective has been to provide the student with clinically relevant questions in a format similar to that encountered on the USMLE Step 1 Boards. The breadth of questions is one of the many features of Lippincott’s Board Review Series titles. We hope that the new edition of BRS Biochemistry, Molecular Biology, and Genetics becomes a valuable tool for students seeking high-yield resources as they prepare for the USMLE Step 1. We recognize the changing nature of science and medicine, however, and encourage readers to send suggestions for improvement for this text or for our companion flash cards, to us via e-mail at LWW.com. Todd Swanson Sandra Kim Marc Glucksman v Publisher’s Preface The Publisher acknowledges the editorial consultation of Michael A. Lieberman, Ph.D., to this fifth edition. In addition to his role as editorial consultant on every chapter, Dr. Lieberman reviewed the entire manuscript to help ensure the accuracy, consis- tency, and timeliness of its content. vii