Harper's Illustrated Biochemistry 26th ed (Lange, 2003) WW.pdf

a LANGE medical book

Illustrated

Biochemistry

twenty-sixth edition

Robert K. Murray, MD, PhD

Professor (Emeritus) of Biochemistry

University of Toronto

Toronto, Ontario

Daryl K. Granner, MD

Joe C. Davis Professor of Biomedical Science

Director, Vanderbilt Diabetes Center

Professor of Molecular Physiology and Biophysics

and of Medicine

Vanderbilt University

Nashville, Tennessee

Peter A. Mayes, PhD, DSc

Emeritus Professor of Veterinary Biochemistry

Royal Veterinary College

University of London

London

Victor W. Rodwell, PhD

Professor of Biochemistry

Purdue University

West Lafayette, Indiana

Lange Medical Books/McGraw-Hill

Medical Publishing Division

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Harper’s

Harper’s Illustrated Biochemistry, Twenty-Sixth Edition

permitted under the United States Copyright Act of 1976, no part of this publication may be reproduced or distributed in any

form or by any means, or stored in a data base or retrieval system, without the prior written permission of the publisher.

234567890DOC/DOC09876543

ISBN 0-07-138901-6

ISSN 1043-9811

Notice

Medicine is an ever-changing science. As new research and clinical experience broaden our knowledge, changes in treat-

ment and drug therapy are required. The authors and the publisher of this work have checked with sources believed to be

reliable in their efforts to provide information that is complete and generally in accord with the standards accepted at the

time of publication. However, in view of the possibility of human error or changes in medical sciences, neither the authors

nor the publisher nor any other party who has been involved in the preparation or publication of this work warrants that

the information contained herein is in every respect accurate or complete, and they disclaim all responsibility for any er-

rors or omissions or for the results obtained from use of the information contained in this work. Readers are encouraged

to confirm the information contained herein with other sources. For example and in particular, readers are advised to

check the product information sheet included in the package of each drug they plan to administer to be certain that the

information contained in this work is accurate and that changes have not been made in the recommended dose or in the

contraindications for administration. This recommendation is of particular importance in connection with new or infre-

quently used drugs.

This book was set in Garamond by Pine Tree Composition

The editors were Janet Foltin, Jim Ransom, and Janene Matragrano Oransky.

The production supervisor was Phil Galea.

The illustration manager was Charissa Baker.

The text designer was Eve Siegel.

The cover designer was Mary McKeon.

The index was prepared by Kathy Pitcoff.

RR Donnelley was printer and binder.

This book is printed on acid-free paper.

ISBN-0-07-121766-5 (International Edition)

manufacture and export. This book cannot be re-exported from the country to which it

is consigned by McGraw-Hill. The International Edition is not available in North America.

Authors

David A. Bender, PhD

Sub-Dean Royal Free and University College Medical

School, Assistant Faculty Tutor and Tutor to Med-

ical Students, Senior Lecturer in Biochemistry, De-

partment of Biochemistry and Molecular Biology,

University College London

Peter A. Mayes, PhD, DSc

Emeritus Professor of Veterinary Biochemistry, Royal

Veterinary College, University of London

Robert K. Murray, MD, PhD

Professor (Emeritus) of Biochemistry, University of

Toronto

Kathleen M. Botham, PhD, DSc

Reader in Biochemistry, Royal Veterinary College,

University of London

Margaret L. Rand, PhD

Scientist, Research Institute, Hospital for Sick Chil-

dren, Toronto, and Associate Professor, Depart-

ments of Laboratory Medicine and Pathobiology

and Department of Biochemistry, University of

Toronto

Daryl K. Granner, MD

Joe C. Davis Professor of Biomedical Science, Director,

Vanderbilt Diabetes Center, Professor of Molecular

Physiology and Biophysics and of Medicine, Vander-

bilt University, Nashville, Tennessee

Victor W. Rodwell, PhD

Professor of Biochemistry, Purdue University, West

Lafayette, Indiana

Frederick W. Keeley, PhD

Associate Director and Senior Scientist, Research Insti-

tute, Hospital for Sick Children, Toronto, and Pro-

fessor, Department of Biochemistry, University of

Toronto

P. Anthony Weil, PhD

Professor of Molecular Physiology and Biophysics,

Vanderbilt University School of Medicine, Nash-

ville, Tennessee

Peter J. Kennelly, PhD

Professor of Biochemistry, Virginia Polytechnic Insti-

tute and State University, Blacksburg, Virginia

vii

Preface

The authors and publisher are pleased to present the twenty-sixth edition of Harper’s Illustrated Biochemistry . Review

of Physiological Chemistry was first published in 1939 and revised in 1944, and it quickly gained a wide readership. In

1951, the third edition appeared with Harold A. Harper, University of California School of Medicine at San Fran-

cisco, as author. Dr. Harper remained the sole author until the ninth edition and co-authored eight subsequent edi-

tions. Peter Mayes and Victor Rodwell have been authors since the tenth edition, Daryl Granner since the twentieth

edition, and Rob Murray since the twenty-first edition. Because of the increasing complexity of biochemical knowl-

edge, they have added co-authors in recent editions.

Fred Keeley and Margaret Rand have each co-authored one chapter with Rob Murray for this and previous edi-

tions. Peter Kennelly joined as a co-author in the twenty-fifth edition, and in the present edition has co-authored

with Victor Rodwell all of the chapters dealing with the structure and function of proteins and enzymes. The follow-

ing additional co-authors are very warmly welcomed in this edition: Kathleen Botham has co-authored, with Peter

Mayes, the chapters on bioenergetics, biologic oxidation, oxidative phosphorylation, and lipid metabolism. David

Bender has co-authored, also with Peter Mayes, the chapters dealing with carbohydrate metabolism, nutrition, diges-

tion, and vitamins and minerals. P. Anthony Weil has co-authored chapters dealing with various aspects of DNA, of

RNA, and of gene expression with Daryl Granner. We are all very grateful to our co-authors for bringing their ex-

pertise and fresh perspectives to the text.

CHANGES IN THE TWENTY-SIXTH EDITION

A major goal of the authors continues to be to provide both medical and other students of the health sciences with a

book that both describes the basics of biochemistry and is user-friendly and interesting. A second major ongoing

goal is to reflect the most significant advances in biochemistry that are important to medicine. However, a third

major goal of this edition was to achieve a substantial reduction in size, as feedback indicated that many readers pre-

fer shorter texts.

To achieve this goal, all of the chapters were rigorously edited, involving their amalgamation, division, or dele-

tion, and many were reduced to approximately one-half to two-thirds of their previous size. This has been effected

without loss of crucial information but with gain in conciseness and clarity.

Despite the reduction in size, there are many new features in the twenty-sixth edition. These include:

•A new chapter on amino acids and peptides, which emphasizes the manner in which the properties of biologic

peptides derive from the individual amino acids of which they are comprised.

•A new chapter on the primary structure of proteins, which provides coverage of both classic and newly emerging

“proteomic” and “genomic” methods for identifying proteins. A new section on the application of mass spectrometry

to the analysis of protein structure has been added, including comments on the identification of covalent modifica-

tions.

• The chapter on the mechanisms of action of enzymes has been revised to provide a comprehensive description of

the various physical mechanisms by which enzymes carry out their catalytic functions.

• The chapters on integration of metabolism, nutrition, digestion and absorption, and vitamins and minerals have

been completely re-written.

• Among important additions to the various chapters on metabolism are the following: update of the information

on oxidative phosphorylation, including a description of the rotary ATP synthase; new insights into the role of

GTP in gluconeogenesis; additional information on the regulation of acetyl-CoA carboxylase; new information on

receptors involved in lipoprotein metabolism and reverse cholesterol transport; discussion of the role of leptin in

fat storage; and new information on bile acid regulation, including the role of the farnesoid X receptor (FXR).

• The chapter on membrane biochemistry in the previous edition has been split into two, yielding two new chapters

on the structure and function of membranes and intracellular traffic and sorting of proteins.

• Considerable new material has been added on RNA synthesis, protein synthesis, gene regulation, and various as-

pects of molecular genetics.

• Much of the material on individual endocrine glands present in the twenty-fifth edition has been replaced with

new chapters dealing with the diversity of the endocrine system, with molecular mechanisms of hormone action,

and with signal transduction.

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