As a new graduate student at Harvard, I heard from experienced teaching fellows that if you were interested in learning how to be an effective teacher, it was essential to seek out extraordinary mentors. Based on word of mouth, I became involved with E. O. Wilson’s course in Evolutionary Biology and Irven DeVore’s course in Human Behavioral Biology. Both were known to be unusually provocative, challenging, and entertaining classes for non-
The two courses were quite different from each other, but at their core both were built on two beliefs that are central to this book and to my thinking about education: (1) Biology is creative, interesting, and fun. (2) Biology is relevant to the daily life of every person. There was a palpable sense that, in teaching non-
My development as a scientist and, particularly, my appreciation for rigorous and methodical critical thinking have been shaped by the kind support and wise guidance of Richard Lewontin. I have also been fortunate to have as a long-
There are many other friends and colleagues I wish to thank for helping me with What Is Life?
In researching and writing the book and in developing the numerous courses I teach, I have benefited from more than a decade of perceptive and valuable contributions, too numerous to list, from Glenn Adelson, Alon Ziv, and Michael Cooperson. I am tremendously appreciative of all they have done for me.
For a project covering so many topics and years, it is essential to have a close group of trusted, tolerant, and knowledgeable colleagues. I am grateful to Alicia Moretti, Harold Owens, Greg Graffin, Brian Swartz, Greg Laden, Jeff Egger, Andy Tobias, Elisabeth Tobias, Joshua Malina, Melissa Merwin-
Numerous colleagues at UCLA provided assistance and support, including Steve Strand, Cliff Brunk, Fred Eiserling, Victoria Sork, Deb Pires, Lianna Johnson, Gaston Pfluegl, Frank Laski, Jeff Thomas, and Tracy Knox. As a result of their commitment to excellence in the UCLA Department of Life Science Core Education, I have been able to acquire a wealth of experiences that have helped me continue improving as a teacher.
I owe a tremendous debt to Sara Tenney, without whose encouragement and support this project could never have been begun or completed.
W. H. Freeman is an extremely author-
It is impossible to teach biology without illustrations. My deepest gratitude goes to Tommy Moorman for creating such innovative and effective figures for the book. Tommy’s vision for an elegant and beautiful art program completely integrated with the text is apparent on every page. Working with him to develop each illustration in this book has been (and continues to be) one of my most enjoyable and satisfying professional collaborations. Thanks also go to Alison Kendall, Tamara Lau, and Erin Daniel for assisting with the creation of the illustrations. For the design of the book, I thank Tom Carling. And for excellent assistance with photo research, thanks to Julia Phelan, Jennifer Atkins, and Christine Buese.
For creating the innovative media and print materials that accompany the book, I am thankful for the contributions of Mike Jones and for the extensive input of Amanda Dunning and Elaine Palucki. I thank all of the contributors and advisors who helped create the student and instructor resources; your efforts have been invaluable. I thank Jennifer Warner and Meredith Norris for their work on the Student Success Guide. I also appreciate the contributions of Troy Williams and the PrepU team. Sheri Snavely provided significant input in developing pedagogical strategies throughout the book; I also appreciate her thoughtful and wise advice at nearly every step in the publishing process. Copyeditor Linda Strange helped to ensure consistency, accuracy, and readability throughout the text. The rest of the life sciences editorial team at W. H. Freeman, too, have been knowledgeable and supportive, particularly Anna Bristow, Lisa Samols, Susan Moran, Kate Parker, and Elaine Palucki. For their efficiency and commitment to producing a beautiful book, I am most grateful to the W. H. Freeman production team: Sheridan Sellers, Elizabeth Geller, Paul Rohloff, Diana Blume, and Catherine Woods.
XXVIII
The people on the marketing team at W. H. Freeman have contributed enormously in helping with the challenging task of introducing this book to students and instructors across the country. John Britch, Shannon Howard, and Todd Elder have been enthusiastic and dedicated in creating materials and strategies to assist instructors in evaluating the ways in which What Is Life? can aid them as they develop their own courses and strategies for success.
Finally, I thank my family—
Contact the author with your feedback.
The content of this book has been greatly improved through the comments of reviewers and students. Your comments, suggestions, and criticism are also welcome; they are essential in guiding its ongoing evolution. Please contact the author at jay@jayphelan.com. I’m serious.
We thank the many reviewers who aided in the development of this text.
Third Edition Development
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XXIX
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Gerald Farr, Texas State University–
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Victor Fet, Marshall University
Steven Fields, Winthrop University
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Mary Beth Finn, Herzing University
Thomas Finnegan, Dunwoody College of Technology
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April Ann Fong, Portland Community College, Sylvania Campus
Donald Fontes, Community College of Rhode Island
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Michael Fox, SUNY–
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Barbara Frank, Idaho State University–
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Kadrin Getman, Blue Ridge Community College
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Philip Gibson, Gwinnett Technical College
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Richard Gill, Brigham Young University
Amy Glaser, Erie Community College
Melissa Glenn, Broome Community College
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Jan Goerrissen, Orange Coast College
Andrew Goliszek, North Carolina A&T State University
Brad Goodbar, College of the Sequoias
Amy Goode, Illinois Central College
Dalton Gossett, Louisiana State University in Shreveport
Tamar Goulet, University of Mississippi
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Jen Grant, University of Wisconsin–
Sherri Graves, Sacramento City College
Madoka Gray-
Melissa Greene, Northwest Mississippi Community College
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John Griffis, Joliet Junior College
Carole Griffiths, Long Island University
Brad Griggs, Piedmont Technical College
Tim Grogan, Valencia College–
Deann Grossi, The Illinois Institute of Art
Laine Gurley, Harper College
Carla Guthridge, Cameron University
Ghislaine Guyot Jackson, Keiser University
Sue Habeck, Tacoma Community College
Nick Hackett, Moraine Valley Community College
Cheryl Hackworth, West Valley College
Mark Haefele, Community College of Denver
William Hairston, Harrisburg Area Community College
Kristy Halverson, University of Southern Mississippi
Michael Hamed, Virginia Highlands Community College
Pamela Hanratty, Indiana University–
Katy Hansen, Bismarck State College
Shari Harden, Metropolitan Community College–
Joyce Hardy, Chadron State College
Janelle Hare, Morehead State University
John Harley, Eastern Kentucky University
Olivia Harriott, Fairfield University
Katherine Harris, Hartnell College
J. Scott Harrison, Georgia Southern University
Jay Hatch, University of Minnesota
Keith Hench, Kirkwood Community College
Krista Henderson, California State University–
Scottie Henderson, Cerritos College
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Gloria Hoffman, Morgan State University
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Amy Hollingsworth, University of Akron
Tina Hopper, Missouri State University
Jane Horlings, Saddleback College
Vanessa Hormann, Broward College
Harold Horn, Lincoln Land Community College
Kris Horn, DeVry University
Anne-
XXX
Laurie Host, Harford Community College
Laura Houston, Northeast Lakeview College
Steve (Robert) Howard, Middle Tennessee State University
Carina Howell, Lock Haven University
Kenny Hudiburg, Fort Scott Community College
Joan Hudson, Sam Houston State University
Vicki Huffman, Potomac State College of West Virginia University
Kimberly Hurd, Bakersfield College
Joseph Husband, Florida State College at Jacksonville
Cynthia Hutton, Northland Pioneer College
Jeba Inbarasu, Metropolitan Community College, South Omaha Campus
Anthony Ippolito, DePaul University
Virginia Irintcheva, Black Hawk College
Kamal Islam, Ball State University
Robert Iwan, Inver Hills Community College
Evelyn Jackson, University of Mississippi
Kristin Jacobson, Illinois Central College
Kathleen Janech, College of Charleston
David Jenkins, University of Alabama at Birmingham
Dianne Jennings, Virginia Commonwealth University
Jamie Jensen, Brigham Young University
Bradley Jett, Oklahoma Baptist University
Carl Johansson, Fresno City College
Mitrick Johns, Northern Illinois University
Keith Johnson, Bradley University
Kelly Johnson, Somerset Community College
Kristy Johnson, The Citadel
Robert Johnson, Pierce College
Staci Johnson, Southern Wesleyan University
Tanganika Johnson, Southern University and A&M College
Gail Jones, Texas Christian University
Gregory Jones, Santa Fe College
Ken Jones, Dyersburg State Community College
Kevin Jones, Charleston Southern University
Jacqueline Jordan, Clayton State University
Glenn Kageyama, California State Polytechnic University, Pomona
Ron Kaltreider, York College of Pennsylvania
Lisa Kaplan, Quinnipiac University
Arnold Karpoff, University of Louisville
Judy Kaufman, Monroe Community College
Jerry Kavouras, Lewis University
Joanna Kazmierczak, Community College of Allegheny County
Dawn Keller, Hawkeye Community College
Kristopher Kelley, University of Louisiana at Monroe
Tonya Kerschner, Butler Community College
Moshe Khurgel, Bridgewater College
Amine Kidane, Columbus State Community College
Michael Kiel, Marywood University
Brandon King, California State Polytechnic University, Pomona
Kevin Kinney, DePauw University
Dennis Kitz, Southern Illinois University, Edwardsville
Mark Knauss, Georgia Highlands College
Jennifer Kneafsey, Tulsa Community College
Nighat Kokan, Cardinal Stritch University
Olga Kopp, Utah Valley University
Mary Korte, Concordia University Wisconsin
Anna Koshy, Houston Community College
Melissa Kosinski-
Karen Koster, University of South Dakota
Peter Kourtev, Carnegie Mellon University
Karen Kowalski, Tidewater Community College
Dennis Kraichely, Cabrini College
Brian Kram, Prince George’s Community College
Rene Kratz, Everett Community College
David Krauss, City University of New York
Joe Kremer, Alvernia University
Tim Kreps, Bridgewater College
Nadine Kriska, University of Wisconsin–
Dana Kurpius, Elgin Community College
Kim Lackey, University of Alabama
Katherine LaCommare, Lansing Community College
Troy Ladine, East Texas Baptist University
Archana Lal, Independence Community College
Kevin Lam, Alexander College
Ellen Lamb, University of North Carolina at Greensboro
Kirkwood Land, University of the Pacific
Gary Lange, Saginaw Valley State University
Patrick Larkin, Santa Fe College
Liz Lawrence, Miles Community College
Tyler Lawson, Baldwin-
Brenda Leady, University of Toledo
Lorraine Leiser, Southeast Community College
Laurie Len, El Camino College
Andrea LeSchack, Trident Technical College
Suzanne Lewis, Truckee Meadows Community College
Harvey Liftin, Broward College
Tammy Liles, Bluegrass Community and Technical College
Debra Linton, Central Michigan University
Kathryn Lipson, Western New England University
Robert Lishak, Auburn University
Cynthia Littlejohn, University of Southern Mississippi
Jason Locklin, Temple College
Suzanne Long, Monroe Community College
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Margaret Lutze, DePaul University
Kimberly Lyle-
Cyrus MacFoy, Montgomery College
William Mackay, Edinboro University of Pennsylvania
Tom Maida, Santa Fe College
Margaret Major, Georgia Perimeter College
Charles Mallery, University of Miami
Cindy Malone, California State University, Northridge
Mark Manteuffel, St. Louis Community College
Lisa Maranto, Prince George’s Community College
Brian Maricle, Fort Hays State University
Marlee Marsh, Columbia College
Paul Marshall, Middlesex Community College
David Martin, Tacoma Community College, Centralia College
Sharon Marusiak, Cuyahoga Community College
Roy Mason, Mt. San Jacinto College
James McCandless, Methodist University
John McDonald, University of Delaware
Tara McGoey, Canadore College
Joan McKearnan, Anoka Ramsey Community College
Karen McLellan, Indiana University–
Margaret McMichael, Baton Rouge Community College
Brock McMillan, Brigham Young University
Daniel Meer, Cardinal Stritch University
Karen Meisch, Austin Peay State University
Sandi Melkonian, Eastern Florida State College
Maryanne Menvielle, California State University, Fullerton
John Mersfelder, Sinclair Community College
Jennifer Metzler, Ball State University
Heather Miceli, Johnson and Wales University
James Mickle, North Carolina State University
Jessica Miles, Palm Beach State College
Craig Milgrim, Grossmont College
Sheila Miracle, Southeast Kentucky Community & Technical College
Jeanne Mitchell, Truman State University
Par Mohammadian, Los Angeles Mission College
Robert Moldenhauer, St. Clair County Community College
Charles Molnar, Camosun College
Thomas Montagno, Mount Wachusett Community College
Scott Moody, Ohio University
Vertigo Moody, Santa Fe College
Brenda Moore, Truman State University
Frances Moore, Patrick Henry Community College
Jeanelle Morgan, University of North Georgia
Erin Morrey, Georgia Perimeter College
Esther Muehlbauer, Queens College
Tim Mulkey, Indiana State University
John Mull, Weber State University
Catherine Murphy, Ocean County College
Christine Mutiti, Georgia College and State University
Elizabeth Nash, Long Beach City College
Rocky Nation, Southern Wesleyan University
Dana Newton, College of The Albemarle
Necia Nicholas, Calhoun Community College
Zia Nisani, Antelope Valley College
Louise Mary Nolan, Middlesex Community College
Meredith Norris, University of North Carolina at Charlotte
Deanna Noyes, Dallas Baptist University
Richard Nuckels, Austin Community College
Judith Ogilvie, St. Louis University
Olumide Ogunmosin, Houston Community College
Katrina Olsen, University of Wisconsin–
Laura O’Riorden, Tallahassee Community College
Joshua Osborn, Henry Ford College
Cassandra Osborne, Western State Colorado University
Mary O’Sullivan, Elgin Community College
Kathleen Page, Bucknell University
David Palmer, Motlow State Community College
Laura Palmer, Penn State Altoona
Joshua Parker, Clayton State University
Ann Paterson, Williams Baptist College
Bruce Patterson, University of Arizona, Tucson
James Peliska, Ave Maria University
Kathleen Pelkki, Saginaw Valley State University
Clayton Penniman, Central Connecticut State University
Marc Perkins, Orange Coast College
Micah Perkins, Owensboro Community and Technical College
Beverly Perry, Houston Community College
Irene Perry, University of Texas of the Permian Basin
John Peters, College of Charleston
Angela Petitclerc, Bishop’s University
Stacy Pfluger, Angelina College
Susan Phillips, Brevard Community College
Don Plantz, Mohave Community College
John Pleasants, Iowa State University
Michael Plotkin, Mt. San Jacinto College
Angela Porta, Kean University
Dan Porter, Amarillo College
Kumkum Prabhakar, Nassau Community College
Saroj Pramanik, Morgan State University
Heather Prestridge, Blinn College
XXXI
Shaun Prince, Lake Region State College
Dorothy Puckett, Kilgore College
Charles Pumpuni, Northern Virginia Community College
Chittur Radhakrishnan, Santa Fe College
Jeffery Ray, University of North Alabama
Mario Raya, MassBay Community College
Samir Raychoudhury, Benedict College
Amy Reber, Georgia State University
Timothy Redman, SUNY Sullivan
Nick Reeves, Mt. San Jacinto College
Rob Reinsvold, University of Northern Colorado
Erin Rempala, San Diego City College
Ashley Rhodes, Kansas State University
Eugenia Ribeiro-
Stanley Rice, Southeastern Oklahoma State University
Brenden Rickards, Gloucester County College
Allison Rober, Ball State University
Leah Roesch, Emory University
Karen Rogowski, Baruch College
Peggy Rolfsen, Cincinnati State Technical and Community College
Chris Romero, Front Range Community College–
Lori Rose, Sam Houston State University
Sadie Rosenthal, Cascadia Community College
Judy Rosovsky, Johnson State College
Hiranya Roychowdhury, New Mexico State University–
Yelena Rudayeva, Palm Beach State College
Lynn Rumfelt, Gordon College
Heather Rushforth, University of North Carolina at Greensboro
Lynette Rushton, South Puget Sound Community College
Michael Rutledge, Middle Tennessee State University
Kim Sadler, Middle Tennessee State University
Stephen Salek, Fayetteville State University
Sydha Salihu, West Virginia University
Shamili Sandiford, College of DuPage
Georgianna Saunders, Missouri State University
Michael Sawey, Texas Christian University
Michael Scanlon, Cornell University
John Schampel, Phoenix Community College
Anna Schmidt, University of Wisconsin–
Emily Schmitt, Nova Southeastern University
Doreen Schroeder, University of St. Thomas
Malcolm Schug, University of North Carolina at Greensboro
Jennifer Scoby, Illinois Central College
Craig Scott, Clarion University
Michael Scott, Lincoln University
Erik Scully, Towson University
Robin Searles-
Pramila Sen, Houston Community College
Carla Serfas, Henry Ford Community College
Brian Seymour, Edward Waters College
Wallace Sharif, Morehouse College
Elizabeth Sharpe-
Dave Sheldon, St. Clair County Community College
Marilyn Shopper, Johnson County Community College
Mark Shotwell, Slippery Rock University
Allan Showalter, Ohio University
Eric Shows, Jones County Junior College
Jack Shurley, Idaho State University
Amanda Simmons, Technical College of the Lowcountry
Jeffrey Simmons, Mount St. Mary’s University
Derek Sims, Hopkinsville Community College
Michael Small, Texas State University
Jennifer Smith, Triton College
Rob Smith, McHenry County College
Tom Smith, Brigham Young University
William Smith, Wake Forest University
Anna Bess Sorin, University of Memphis
Kathryn Sparace, Tri-
Salvatore Sparace, Clemson University
Ashley Spring, Brevard Community College; Eastern Florida State College
William Sproat, Walters State Community College
Brooke Stabler, University of Central Oklahoma
Sonja Stampfler, Kellogg Community College
Sharon Standridge, Middle Georgia State College
Wendy Stankovich, University of Wisconsin–
John Starnes, Somerset Community College
Nancy Staub, Gonzaga University
Thomas Stege, Austin Community College
Mikel Stevens, Brigham Young University
Jon Stoltzfus, Michigan State University
Lisa Strong, Northwest Mississippi Community College
Irina Stroup, Shasta College
Julie Stupi, Lakeland Community College
Aaron Sullivan, Houghton College
Susan Sullivan, Louisiana State University at Alexandria
Qiang Sun, University of Wisconsin–
Kirby Swenson, Middle Georgia College
Joyce Tamashiro, University of Puget Sound
David Tapley, Salem State University
Tatiana Tatum Parker, St. Xavier University
Kimberly Taugher, Diablo Valley College
Mike Taylor, Santiago Canyon College
Shervia Taylor, Southern University
Franklyn Te, Miami Dade College
Don Terpening, Ulster County Community College
Pamela Thinesen, Century College
Jeffrey Thomas, Queens University of Charlotte
Anna Thompson, Feather River College
Jamey Thompson, Hudson Valley Community College
Rita Thrasher, Pensacola State College
Nina Thumser, Edinboro University of Pennsylvania
Candace Timpte, Georgia Gwinnett College
Jonathan Titus, SUNY Fredonia
Anne Tokazewski, Burlington County College
Robert Tompkins, Belmont Abbey College
Willetta Toole-
Michelle Tremblay, Georgia Southern University
Dan Trubovitz, San Diego Miramar College
Anh-
Marsha Turell, Houston Community College
Corey Turner, Lone Star College
Muatasem Ubeidat, Southwestern Oklahoma State University
Sophia Ushinsky, Concordia University
Jagan Valluri, Marshall University
Tom Vance, Navarro College
Pete van Dyke, Walla Walla Community College
Eric Vaught, Northwest Arkansas Community College
Jose Vazquez, New York University
Leticia Vega, Barry University
Aggie Veld, Olivet Nazarene University
William Velhagen, New York University
Uma Vithala, Des Moines Area Community College
Jennifer Vlk, Elgin Community College
Ann Vogelmann, Augustana College
Amie Voorhees, Fresno City College
Walt Wagner, Rowan University
Katherine Warpeha, University of Illinois Chicago
Kathy Webb, Bucks County Community College
Colby Weeks, Brigham Young University–
Charles Welsh, Duquesne University
Michael Wenzel, Folsom Lake College
Alexander Werth, Hampden-
Alicia Whatley, Troy University
Clay White, Lone Star College–
Jennifer Wiatrowski, Pasco-
Rachel Wiechman, West Liberty University
Torelen Winbush, Westwood College
Donnell Young, Waubonsee Community College
John Zamora, Middle Tennessee State University
Lara Zerkowski, Albright College
Ted Zerucha, Appalachian State University
Brenda Zink, Northeastern Junior College
Contributors and Advisors
Instructor’s Manual—
Verona A. Barr, Heartland Community College
Stephen M. Gómez, Central New Mexico Community College
Paul H. Marshall, Northern Essex Community College
Exploring Biology: Case Studies
Michelle Cawthorn, Georgia Southern University
Jennifer L. Holzman, Emory University
Figure Conversion Engine
Ann Aguanno, Marymount Manhattan College
Hands-
Mimi Bres, Cassandra Moore-
Image Bank Advisor
Jane J. Henry, Baton Rouge Community College
Keynote Lecture Presentation
Michael C. Bucher, College of San Mateo
PowerPoint Lecture Outlines
Kristen L. Curran, University of Wisconsin–
Danielle DuCharme, Waubonsee Community College
Jennifer Lange, Chabot College
Mark S. Manteuffel, St. Louis Community College
Prep-
Glenn Adelson, Lake Forrest College
Jay Phelan, University of California, Los Angeles
Alon Ziv, Prep-
Student Worksheets
Michael C. Bucher, College of San Mateo
Active Learning Lecture Outlines
Kristen L. Curran, University of Wisconsin-
Mark S. Manteuffel, St. Louis Community College
Bio 101 Tutorials
Amy Horner-
Kavita Oommen, Georgia State University
Biology Connections Articles
Amy Coombs
Paul H. Marshall, Northern Essex Community College