Fifty years ago, Al Lehninger published the first edition of Biochemistry, defining the basic shape of biochemistry courses worldwide for generations. We are honored to have been able to carry on the Lehninger tradition since his passing in 1986, now introducing the eighth (our seventh) edition of Lehninger Principles of Biochemistry.
This book is a team effort, and producing it would be impossible without the outstanding people at Macmillan Learning who have supported us at every step along the way. Elizabeth Simmons, Program Manager, Biochemistry, led us fearlessly into the brave new world of textbook publishing in the media age. Catherine Murphy, Development Editor, helped develop the revision plan for this edition, cheerfully kept us focused on that plan, skillfully evaluated reviewer comments, and edited the text with a clear eye. Vivien Weiss, Senior Content Project Manager, put all the pieces together seamlessly. Diana Blume, Natasha Wolfe, Maureen McCutcheon, and John Callahan are responsible for the vibrant design of the text and cover of the book. Adam Steinberg and Emiko Paul created the new art for this edition. Photo Researcher Jennifer Atkins and Media Permissions Manager Christine Buese located images and obtained permission to use them. Cate Dapron copy edited and Paula Pyburn proofread the text. Karen Misler, Editorial Project Manager, and Senior Workflow Project Manager Paul W. Rohloff worked diligently to keep us on schedule, and Nathan Livingston helped orchestrate reviews and provided administrative assistance. Cassandra Korsvik and Kelsey Hughes, Media Editors, and Jim Zubricky, Learning Solutions Specialist, oversaw the enormous task of creating the many interactive media enhancements of our content. Our gratitude also goes to Maureen Rachford, Senior Marketing Manager, for coordinating the sales and marketing efforts that bring Lehninger Principles of Biochemistry to the attention of teachers and learners.
In Madison, Brook Soltvedt is, and has been for all the editions we have worked on, our invaluable first-line editor and critic. She is the first to see manuscript chapters, aids in manuscript and art development, ensures internal consistency in content and nomenclature, and keeps us on task with more-or-less gentle prodding. Much of the art and molecular graphics was created by Adam Steinberg of Art for Science, who often made valuable suggestions that led to better and clearer illustrations. The deft hand of Linda Strange, who copyedited six editions of this textbook (including the first), is still evident in the clarity of the text. We feel very fortunate to have had such gifted partners as Brook, Adam, and Linda on our team. We are also indebted to Brian White of the University of Massachusetts Boston, who wrote most of the data analysis problems at the end of chapters.
Many others helped us shape this eighth edition with their comments, suggestions, and criticisms. To all of them, we are deeply grateful:
Ravinder Abrol, California State University–Northridge
Paul Adams, University of Arkansas
Balasubrahmanyam Addepalli, University of Cincinnati
Richard Amasino, University of Wisconsin–Madison
Darryl Aucoin, Caldwell College
Gerald F. Audette, York University, North York
Amy Babbes, Scripps College
Kenneth Balazovich, University of Michigan
Sandra Barnes, Alcorn State University
David Bartley, Adrian College
Zeenat Bashir, Canisius College
Dana Baum, St. Louis University–Main Campus
Donald Beitz, Iowa State University
Henrike Besche, Harvard Medical School
Mrinal Bhattacharjee, Long Island University–Brooklyn
Joshua M. Blose, College at Brockport–State University of New York
Paul Bond, Shorter University
Michael Borenstein, Temple University School of Pharmacy
Kevin Brown, University of Florida–Gainesville
Robert Brown, Memorial University of Newfoundland
D. Andrew Burden, Middle Tennessee State University
Christopher Cottingham, University of North Alabama
Garland Crawford, Mercer University–Macon
Tanya Dahms, University of Regina
Tuhin Das, John Jay College of Criminal Justice, CUNY
Susan Colette Daubner, St. Mary’s University
Margaret Daugherty, Colorado College
Paul DeLaLuz, Lee University
Natasha DeVore, Missouri State University Springfield
Justin DiAngelo, Pennsylvania State University–Berks Campus
Tomas T. Ding, North Carolina Central University
Kristin Dittenhafer-Reed, Hope College
Cassidy Dobson, Truman State University
Artem Domashevskiy, John Jay College of Criminal Justice, CUNY
Donald Doyle, Georgia Institute of Technology
David H. Eagerton, Campbell University
Daniel Edwards, California State University–Chico
Steven Ellis, University of Louisville
Chandrakanth Emani, Western Kentucky University–Bowling Green
Nuran Ercal, Missouri University of Science & Technology
Stylianos Fakas, Alabama A & M University
Russ Feirer, St. Norbert College & Medical College of Wisconsin
Kirsten Fertuck, Northeastern University–Boston
Jennifer Fishovitz, Saint Mary’s College
Kathleen Foley Geiger, Michigan State University
Marcello Forconi, College of Charleston
Isaac Forquer, Portland State University
Jason Fowler, Lincoln Memorial University
Kevin Francis, Texas A&M–Kingsville
Jean Gaffney, Baruch College
Katie Garber, St. Norbert College
Ronald Gary, University Nevada–Las Vegas
Yulia Gerasimova, University of Central Florida
Dipak K. Ghosh, North Carolina A & T State University
Marina Gimpelev, Dominican College
Burt Goldberg, New York University
Daniel Golemboski, Bellarmine College–Louisville
Lawrence Gracz, Massachusetts College of Pharmacy & Health Sciences
Jennifer E. Grant, University of Wisconsin Stout
Joel Gray, Texas State University
Amy Greene, Albright College
Nicholas Grossoehme, Winthrop University
Neena Grover, Colorado College
Rishab K. Gupta, University of California–Los Angeles
Paul Hager, East Carolina University
Bonnie Hall, Grand View University
Marilena Hall, Stonehill College
Christopher Hamilton, Hillsdale College
Matthew Hartman, Virginia Commonwealth University
Mary Hatcher-Skeers, Scripps College
Robin Haynes, Harvard University
Tamara Hendrickson, Wayne State University
Newton Hilliard, Arkansas Technical University
Danny Ho, Columbia University–New York
Jane Hobson, Kwantlen Polytechnic University
Charles Hoogstraten, Michigan State University
Amber Howerton, Nevada State College
Tom Huxford, San Diego State University
Cheryl Ingram-Smith, Clemson University
Lori Isom, University of Central Arkansas
Bruce Jacobson, St. Cloud State University
Nitin Jain, University of Tennessee
Blythe Janowiak, St. Louis University
Matthew R. Jensen, Concordia University, St. Paul
Joseph Jez, Washington University in St. Louis
Xiangshu Jin, Michigan State University–East Lansing
Gerwald Jogl, Brown University
Todd Johnson, Weber State University–Ogden
Marjorie A. Jones, Illinois State University
P. Matthew Joyner, Pepperdine University
Christopher Jurgenson, Delta State University
Jason Kahn, University of Maryland
Kalju Kahn, University of California–Santa Barbara
Peter Kahn, Rutgers University
Reza Karimi, Pacific University
Bhuvana Katkere, Pennsylvania State University–Main Campus
Kevin Kearney, MCPHS University
Chu-Young Kim, University of Texas–El Paso
Bryan Knuckley, University of North Florida
Michael Koelle, Yale University
Andy Koppisch, Northern Arizona University
Joanna Krueger, University of North Carolina–Charlotte
Terry Kubiseski, York University–Keele Campus
Maria Kuhn, Madonna University
Chandrika Kulatilleke, City University of New York–Baruch College
Allison Lamanna, University of Michigan–Ann Arbor
Kimberly Lane, Radford University
Patrick Larkin, Texas A&M University–Corpus Christi
Paul Larsen, University of California–Riverside
Heather Larson, Indiana University Southeast
Benjamin Lasseter, Christopher Newport University
Katherine Launer-Felty, Connecticut College
James Lee, Old Dominion University
Sarah Lee, Abilene Christian University
Scott Lefler, Arizona State University–Tempe
Pingwei Li, Texas A&M University
Yingchun Li, Texas A&M University–Prairie View
Yun Li, Delaware Valley University
Andy LiWang, University of California–Merced
Kimberly Lyle-Ippolito, Anderson University
Taylor J. Mach, Concordia University, St. Paul
Meagan Mann, Austin Peay State University
Glover Martin, University of Massachusetts–Boston
Michael Massiah, George Washington University
Brannon McCullough, Northern Arizona University
John Means, University of Rio Grande
Michael Mendenhall, University of Kentucky
Sabeeha Merchant, University of California–Berkeley
Elizabeth Middleton, Purchase College, SUNY
Jeremy T. Mitchell-Koch, Bethel College–North Newton
Somdeb Mitra, New York University
Susan Mitroka, Worcester State University
Judy Moore, Lenoir-Rhyne University–Hickory
Graham Moran, Loyola University Chicago
Fares Najar, University of Oklahoma–Norman
Scott Napper, University of Saskatchewan
Allen Nicholson, Temple University–Philadelphia
James Nolan, Georgia Gwinnett College
George Nora, Northern State University
Grazyna Nowak, University of Arkansas for Medical Sciences
Abdel Omri, Laurentian University
Allyn Ontko, Arkansas State University
Siva Panda, Augusta State University
Amanda Parker, William Cary University
Jonathan Parrish, University of Alberta
Donna Pattison, University of Houston
Craig Peebles, University of Pittsburgh
Mary Elizabeth Peek, Georgia Institute of Technology Main Campus
Mario Pennella, University of Wisconsin–Madison
Michael Pikaart, Hope College
Deborah Polayes, George Mason University
Alfred Ponticelli, University at Buffalo, Jacobs School of Medicine and Biomedical Sciences
Tamiko Porter, Indiana University–Purdue University Indianapolis
Michelle Pozzi, Texas A&M University
Ramin Radfar, Wofford College
Kevin Redding, Arizona State University
Tanea Reed, Eastern Kentucky University
Christopher Reid, Bryant University
John Richardson, Austin College
Katarzyna Roberts, Rogers State University
Jim Roesser, Virginia Commonwealth University
Christopher Rohlman, Albion College
Brenda Royals, Park University
Gillian Rudd, Georgia Gwinnett College
Megan E. Rudock, Wake Forest University
Joshua Sakon, University of Arkansas–Fayetteville
Nianli Sang, Drexel University
Pradip Sarkar, Parker University
Patrick Schacht, California Baptist University
Kersten Schroeder, University of Central Florida
Joseph Schulz, Occidental College
Michael Sehorn, Clemson University
Kavita Shah, Purdue University–Main Campus
Wallace Sharif, Morehouse College
Ike Shibley, Pennsylvania State University–Berks Campus
Aaron Sholders, Colorado State University Fort Collins
Kevin Siebenlist, Marquette University
Richard Singiser, Clayton State University
Corin Slown, California State University Monterey Bay
Kelli Slunt, University of Mary Washington
Kerry Smith, Clemson University
Sarah J. Smith, Bucknell University
Mark Snider, The College of Wooster
Jennifer Sniegowski, Arizona State University–Downtown
David Snyder, William Paterson University
Joshua Sokoloski, Salisbury University
Joanne Souza, Stony Brook University
Narasimha Sreerama, Colorado State University
Rekha Srinivasan, Case Western Reserve University
Aleksandra Stamenov, University of California–Merced
Ryan Steed, University of North Carolina–Asheville
Angela K. Stoeckman, Bethel University
Koni Stone, California State University Stanislaus
Evelyn Swain, Presbyterian College
Jeremy Thorner, University of California–Berkeley
Candace Timpte, Georgia Gwinnett College
Jamie Towle-Weicksel, Rhode Island College
Michael Trakselis, Baylor University
Brian Trewyn, Colorado School of Mines
Vishwa D. Trivedi, Bethune Cookman University
Didem Vardar-Ulu, Boston University
Thomas Vida, University of Houston
Lori Wallrath, University of Iowa–Iowa City
Chris Wang, Ambrose University
Xuemin Wang, University of Missouri–St. Louis
Yu Wang, The University of Alabama
Robert J. Warburton, Shepherd University
Todd M. Weaver, University of Wisconsin La Crosse
James D. West, The College of Wooster
Chuan Xiao, University of Texas–El Paso
Alexander G. Zestos, American University
We lack the space here to acknowledge all the other individuals whose special efforts went into this book. We offer instead our sincere thanks—and the finished book that they helped guide to completion. We, of course, assume full responsibility for errors of fact or emphasis.
We want especially to thank our students at the University of Wisconsin–Madison for their numerous comments and suggestions. If something in the book does not work, they are never shy about letting us know it. We are grateful to the students and staff of our past and present research groups, who helped us balance the competing demands on our time; to our colleagues in the Department of Biochemistry at the University of Wisconsin–Madison, who helped us with advice and criticism; and to the many students and teachers who have written to suggest ways of improving the book. We hope our readers will continue to provide input for future editions.
Finally, we express our deepest appreciation to our partners (Brook, Beth, and Tim) and our families, who showed extraordinary patience with, and support for, our book writing.