LEHNINGER PRINCIPLES OF BIOCHEMISTRY - VOL. 1. THE FOUNDATIONS OF BIOCHEMISTRY: STRUCTURE AND CATALYSIS - 2011

Preface

The first edition of Principles of Biochemistry, written by Albert Lehninger 25 years ago, served as a foundational starting point and model for the four subsequent editions. Over the past quarter-century, biochemical knowledge has expanded exponentially. Twenty-five years ago, not a single genome had been sequenced, no X-ray crystallographic Structure of a membrane protein had been solved, and knock-out mice did not yet exist. Ribozymes and RT-PCR technology had only just been introduced, and Archaea were just beginning to be recognized as a distinct domain of life. Today, new genetic sequences are published weekly, and novel protein structures emerge even more frequently; researchers have generated thousands of different knock-out animal models, driving remarkable advancements in biochemistry, physiology, and medicine. This fifth edition features photographs of 31 Nobel laureates who received their prizes in chemistry, physiology, or medicine after the publication of the very first edition.

A constant challenge for each new edition has been to capture the flood of incoming information without overwhelming students, many of whom are encountering biochemistry for the very first time. Rigorous Selection has allowed us to present the core principles of biochemistry while highlighting exciting recent scientific breakthroughs and their implications for humanity's future. A prime example of such groundbreaking research is the X-ray crystallographic analysis of RNA polymerase, which has made it possible to visualize, at atomic resolution, the informational roles of DNA, RNA, and protein at the precise moment of information transfer.

We stand on the threshold of a new molecular physiology—an era in which processes such as membrane excitation, secretion, hormone action, Vision, Olfaction, taste, Respiration, Muscle contraction, and Cell motility will be explained at THE MOLECULAR LEVEL and made fully accessible to genetic analysis and pharmacological intervention. Understanding the molecular architecture of complex membrane systems, such as those responsible for Oxidative Phosphorylation AND Photophosphorylation, will undoubtedly deepen our insight into these vital life processes. (Such rapid progress almost makes us wish we could be young again and just starting our academic and teaching careers—after all, our textbook isn't the only thing that has aged over the years.)

In every new edition, we have strived to preserve the defining qualities that made Lehninger a classic textbook: clarity of exposition, meticulous explanation of challenging concepts, and a strong emphasis on the experimental Methods used by modern biochemists. As authors, we have worked together for over 25 years, and nearly as long as educators. Throughout this time, thousands of our students at the University of Wisconsin–Madison have been an endless source of inspiration on how to make The Study of modern biochemistry more accessible. Our students have taught and inspired us. It is our sincere hope that this twenty-fifth-anniversary edition will similarly educate and inspire a new generation of biochemistry students, perhaps even instilling in some of them the same enduring passion for the subject that we share.

Class="center">Acknowledgments

This book is a collaborative endeavor, and its publication would not have been possible without the unwavering support of the W. H. Freeman and Company team who guided us every step of the way. Randy Rossiroli (Editor-in-Chief) and Kate Ahr (Managing Editor) organized reviews, offered countless invaluable suggestions, encouraged us, kept us on track, and heroically (though not always successfully) reined us in within our deadlines. Our outstanding developmental editor, Liz Heller, somehow managed to shepherd the book through production despite missed deadlines and sudden revisions, always doing so with her characteristic charm and expertise. We are grateful to Vicky Tomaselli for the book design and Marsha Cohen for the magnificent layout. We were once again fortunate to work with our brilliant senior manuscript editor, Linda Strange, who has copyedited all five editions of Principles of Biochemistry (as well as Lehninger’s two early editions of Biochemistry). Her assistance is immeasurable, and every Touch she gives to the text unfailingly improves it. We are also delighted to have worked alongside Morgan Ryan, who assisted us with the previous two editions. We thank Dena Digilio Betz for her tireless work in sourcing and coordinating photographs, and Nick Timoshko and Whitney Clench for keeping all the moving PARTS OF THE project synchronized. Finally, we express our gratitude to Debbie Clare, Director of Marketing, for her exceptional sales and marketing efforts and her wonderful sense of humor.

In Madison, our primary in-house editor and critic throughout all editions has been, and remains, Brooke Saltvedt. She is invariably the first to read through manuscript chapters, assisting with text and illustration preparation, checking cross-References, ensuring consistent nomenclature, and gently yet persistently prodding us forward. We also thank Shelly Luzzetti, now at the University of New Mexico, who, just as for the previous edition, proofread every single word of the text, caught numerous errors, and provided astute suggestions that greatly enhanced the book.

The fresh visual design of the book, including the rendering of new molecular structures, was crafted right here in Madison by Adam Steinberg, whose insightful suggestions made the illustrations vastly more communicative. This edition also features molecular art created for the third and fourth editions by our Madison colleague Jean-Yves Sgro. We consider ourselves immensely fortunate to collaborate with such exceptionally talented individuals as Brooke, Shelly, Adam, and Jean-Yves.

We are likewise deeply indebted to Brian White of the University of Massachusetts Boston, who developed the new data-analysis problem sets for each chapter of the book.

Numerous colleagues followed the progress of this project with keen interest and provided invaluable assistance through their thoughtful comments. We wish to express our special gratitude to Lawrence Anderson of the University of Wisconsin–Madison, Jeffrey D. Esko of the University of California, San Diego, Jack Kirsch and his students at the University of California, Berkeley, as well as Dana Ashworth, Shiow-Chuall (Cheryl) Tsai, Michael G. Cumsky, and their colleagues (listed below) from the University of California, Irvine. Many individuals contributed to the shape and execution of this fifth edition with their expert advice, reviews, and critiques. We gratefully acknowledge everyone included in the following list:

Richard M. Amasino, University of Wisconsin–Madison

Louise E. Anderson, University of Illinois at Chicago

Cheryl Bailey, University of Nebraska–Lincoln

Kenneth Balazovits, University of Michigan

Thomas O. Baldwin, University of Arizona

Vaji Bhandarian, University of Arizona

Eugene Barber, University of Rochester

Sebastian I. Bednarek, University of Wisconsin–Madison

Ramachandra Bhat, Lincoln University

James Blankenship, Cornell University

Sandra J. Bonetti, Colorado State University–Pueblo

Barbara Bowman, University of California, Berkeley

Scott D. Briggs, Purdue University

Jeff Brodsky, University of Pittsburgh

Ben Caldwell, Western Missouri State University

David Camerini, University of California, Irvine

Jeffrey Kohlberg, California State University, Long Beach

Guillaume Chanfreau, University of California, Los Angeles

Melanie Cocco, University of California, Irvine

Kim D. Collins, University of Maryland

Charles T. Dameron, Duquesne University

Richard S. Eisenstein, University of Wisconsin–Madison

Gerald W. Feigenson, Cornell University

Robert G. Fillingame, University of Wisconsin–Madison

Brian Fox, University of Wisconsin–Madison

Gerald D. Frenkel, Rutgers University

Perry Frey, University of Wisconsin–Madison

David E. Graham, University of Texas at Austin

William J. Grimes, University of Arizona

Martin Gunn, Texas A&M University

Olivia Hanson, University of Central Oklahoma

Amy Hark, Muhlenberg College

Sean W. Hernandez, University of Wisconsin–Madison

Peter Hinkle, Cornell University

P. Shing Ho, University of Oregon

Charles G. Hoogstraten, Michigan State University

Gerwald Jogl, Brown University

Sir Hans Kornberg, Boston University

Bob Landick, University of Wisconsin–Madison

Patrick D. Larkin, Texas A&M University–Corpus Christi

Ryan P. Liegel, University of Wisconsin–Madison

Maria Linder, California State University, Fullerton

Andy S. LiWang, Texas A&M University

John Makemson, Florida International University

John S. Matthews, University of Mississippi School of Pharmacy

Benjamin J. McFarland, Seattle Pacific University

Anant Menon, Weill Cornell Medical College

Sabeeha Merchant, University of California, Los Angeles

Scott S. Mohr, Boston University

Kimberly Mowry, Brown University

Leisha Mullins, Texas A&M University

Sewayéh Negash, California State University, Long Beach

Allen W. Nicholson, Temple University

Hiroshi Nikaido, University of California, Berkeley

James Ntambi, University of Wisconsin–Madison

Timothy F. Osborne, University of California, Irvine

José R. Pérez-Castejón, University of Seville, Spain

Terry Platt, University of Rochester

Wendy Pogozelski, State University of New York at Geneseo

Jonathan Popper, University of Wisconsin–Madison

Thomas Paulos, University of Wisconsin–Madison

Jack Preiss, Michigan State University

Anna Rodaminsk-Pandia, University of Arkansas

Ron Raines, University of Wisconsin–Madison

Tom A. Rapoport, Harvard Medical School

Jason J. Reddick, University of North Carolina,

Greensboro Mary Roberts, Boston College Ingrid

C. Ruth, University of California, Irvine

Abuzar Soleimani, University of Tehran, Iran

Mark Spaller, Wayne State University

Stephen Spiro, University of Texas at Dallas

Narasimha Srirama, University of Colorado Ion

D. Stewart, University of Florida

Connie Stone, California State University, Stanislaus

Ion R. Stultsfoos, University of Michigan

Jeremy Thorner, University of California, Berkeley

Dean R. Tolan, Boston University

Sandra L. Turchi, Millersville University

Manuel Varela, Eastern New Mexico University

Bob Warburton, Shepherd University

Tracey Ware, Salem College

Susan Weintraub, University of Texas Health Science Center

Michael Jaffe, Massachusetts Institute of Technology

We simply do not have enough space to thank everyone whose help we relied upon during the preparation of this book for publication. We express our deepest gratitude to all of them, and this completed work serves as a testament to our appreciation. Naturally, we assume full responsibility for any errors or inaccuracies that may still be found in the book.

We would like to thank our students at the University of Wisconsin–Madison once again for their numerous suggestions and comments. Should anything be amiss in the book, they will undoubtedly let us know. We are grateful to the students and members of our research groups, as well as the staff of the Center for Biology Education, who helped us manage our time effectively; we thank our colleagues in the Department of Biochemistry at the University of Wisconsin–Madison for their advice and critical feedback; and we extend our appreciation to all the students and instructors who have sent us their comments and suggestions. We hope our readers will continue to contribute to future editions of this book.

Finally, we express our utmost gratitude to our wives, Brook and Beth, and to our families, whose remarkable patience and unwavering support made the writing of this book possible.

David L. Nelson, Michael M. Cox, Madison, Wisconsin, January 2008



Last update: 06/08/2026

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