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This is an authoritative introductory text that presents biological concepts through the research that revealed them. "Life" covers the full range of topics with an integrated experimental focus that flows naturally from the narrative.
Biology is a constantly changing scientific field. New discoveries about the living world are being made every day, and more than 1 million new research articles in biology are published each year. Beyond the constant need to update the concepts and facts presented in any science textbook, in recent years ideas about how best to educate the upcoming generation of biologists have undergone dynamic and exciting change. Although we and many of our colleagues had thought about the nature of biological education as individuals, it is only recently that biologists have come together to discuss these issues. Reports from the National Academy of Sciences, Howard Hughes Medical Institute, and College Board AP Biology Program not only express concern about how best to instruct undergraduates in biology, but offer concrete suggestions about how to design the introductory biology course—and by extension, our book. We have followed these discussions closely and have been especially impressed with the report “Vision and Change in Undergraduate Biology Education” (visionandchange. org). As participants in the educational enterprise, we have answered the report’s call to action with this textbook and its associated ancillary materials. The “Vision and Change” report proposes five core concepts for biological literacy: 1. Evolution 2. Structure and function 3. Information flow, exchange, and storage 4. Pathways and transformations of energy and matter 5. Systems These five concepts have always been recurring themes in Life, but in this Tenth Edition we have brought them even more “front and center.” “Vision and Change” also advocates that students learn and demonstrate core competencies, including the ability to apply the process of science using quantitative reasoning. Life has always emphasized the experimental nature of biology. This edition responds further to these core competency issues with a new working with data feature and the addition of a statistics primer (Appendix B). The authors’ multiple educational perspectives and areas of expertise, as well as input from many colleagues and students who used previous editions, have informed the approach to this new edition. Enduring Features We remain committed to blending the presentation of core ideas with an emphasis on introducing students to the process of scientific inquiry. Having pioneered the idea of depicting important experiments in unique figures designed to help students understand and appreciate the way scientific investigations work, we continue to develop this approach in the book’s 70 Investigating Life figures. Each of these figures sets the experiment in perspective and relates it to the accompanying text. As in previous editions, these figures employ the structure Hypothesis, Method, Results, and Conclusion. We have added new information focusing on the individuals who performed these experiments so students can appreciate more fully that science is a human and very personal activity. Each Investigating Life figure has a reference to BioPortal (, where discussion and references to follow-up research can be found. A related feature is the Research Tools figures, which depict laboratory and field methods used in biology. These, too, have been expanded to provide more useful context for their importance. Some 15 years ago, Life’s authors and publishers pioneered the use of balloon captions in our figures. We recognized then that many students are visual learners, and this fact is even truer today. Life’s balloon captions bring the crucial explanations of intricate, complex processes directly into the illustration, allowing students to integrate information without repeatedly going back and forth between the figure, its legend, and the text. We continue to refine our chapter organization. Our opening stories have always provide historical, medical, or social context to intrigue students and show how the subject of each chapter relates to the world around them. In the Tenth Edition, the opening stories all end with a question that is revisited throughout the chapter. At the end of each chapter the answer is presented in the light of material the student encountered in the body of the chapter. A chapter outline asks questions to emphasize scientific inquiry, each of which is answered in a major section of the chapter. A Recap summarizes each section’s key concepts and poses questions that help the student review and test their mastery of these concepts. The recap questions are similar in form to the learning objectives used in many introductory biology courses. The Chapter Summaries highlight each chapter’s key figures and defined terms, while restating the major concepts presented in the chapter in a concise and student-friendly manner, with references to specific figures and to the activities and animated tutorials available in BioPortal. At the end of the book, students will find a much-expanded glossary that continues Life’s practice of providing Latin or Greek derivations for many of the defined terms. As students become gradually (and painlessly) more familiar with such root words, the mastery of vocabulary as they continue in their biological or medical studies will be easier. In addition, the popular Tree of Life appendix (Appendix A) presents the phylogenetic tree of life as a reference tool that allows students to place any group of organisms mentioned in the text into the context of the rest of life. The web-based version of Appendix A provides links to photos, keys, species lists, distribution maps, and other information (via the online database at to help students explore biodiversity in greater detail. New Features The Tenth Edition of Life has a different look and feel from its predecessors. The new color palette and more open design will, we hope, be more accessible to students. And, in keeping with our heightened emphasis on scientific inquiry and quantitative analysis, we have added Working with Data exercises to almost all chapters. In these innovative exercises, we describe the context and approach of a research paper that provides the basis of the analysis. We then ask questions that require students to analyze data, make calculations, and draw conclusions. Answers (or suggested possible answers) to these questions are included in BioPortal and can be made available to students at the instructor’s discretion. Because many of the questions in the Working with Data exercises require the use of basic statistical methods, we have included a Statistics Primer as the book’s Appendix B, describing the concepts and some methods of statistical analysis. We hope that the Working with Data exercises and statistics primer will reinforce students’ skills and their ability to apply quantitative analysis to biology. We have added links to Media Clips in the body of the text, with at least one per chapter. These brief clips are intended to enlighten and entertain. Recognizing the widespread use of “smart phones” by students, the textbook includes instant access (QR) codes that bring the Media Clips, Animated Tutorials, and Interactive Summaries directly to the screen in your hand. If you do not have a smart phone, never fear, we also provide direct web addresses to these features. As educators, we follow current discussions of pedagogy in biological education. The chapter-ending Chapter Reviews now contain multiple levels of questions based on Bloom’s taxonomy: Remembering, Understanding and Applying, and Analyzing and Evaluating. Answers to these questions appear at the end of the book. For a detailed description of the media and supplements available for the Tenth Edition, please turn to “Life’s Media and Supplements” on page xvii. The Ten Parts PART ONE, THE SCIENCE OF LIFE AND ITS CHEMICAL BASIS Chapter 1 introduces the core concepts set forth in the “Vision and Change” report and continues the much-praised approach of focusing on a specific series of experiments that introduces students to biology as an experimentally based and constantly expanding science. Chapter 1 emphasizes the principles of biology that are the foundation for the rest of the book, including the unity of life at the cellular level and how evolution unites the living world. Chapters 2–4 cover the chemical principles and building blocks that underlie life. Chapter 4 also includes a discussion of how life could have evolved from inanimate chemicals. PART TWO, CELLS The nature of cells and their role as the structural and functional basis of life is foundational to biology. These revised chapters include expanded explanations of how experimental manipulations of living systems have been used to discover cause and effect in biology. Students who are intrigued by the question “Where did the first cells come from?” will appreciate the updated discussion of ideas on the origin of cells and organelles, as well as expanded discussion of the evolution of multicellularity and cell interactions. In response to reviewer comments, the discussion of membrane potential has been moved to Chapter 45, where students may find it to be more relevant. PART THREE, CELLS AND ENERGY The biochemistry of life and energy transformations are among the most challenging topics for many students. We have worked to clarify such concepts as enzyme inhibition, allosteric enzymes, and the integration of biochemical systems. Revised presentations of glycolysis and the citric acid cycle now focus, in both text and figures, on key concepts and attempt to limit excessive detail. There are also revised discussions of the ecological roles of alternate pathways of photosynthetic carbon fixation, as well as the roles of accessory pigments and reaction center in photosynthesis. PART FOUR, GENES AND HEREDITY This crucial section of the book is revised to improve clarity, link related concepts, and provide updates from recent research results. Rather than being segregated into separate chapters, material on prokaryotic genetics and molecular medicine are now interwoven into relevant chapters. Chapter 11 on the cell cycle includes a new discussion of how the mechanisms of cell division are altered in cancer cells. Chapter 12 on transmission genetics now includes coverage of this phenomenon in prokaryotes. Chapters 13 and 14 cover gene expression and gene regulation, including new discoveries about the roles of RNA and an expanded discussion of epigenetics. Chapter 15 covers the subject of gene mutations and describes updated applications of medical genetics. PART FIVE, GENOMES This extensive and up-to-date coverage of genomes expands and reinforces the concepts covered in Part Four. The first chapter of Part Five describes how genomes are analyzed and what they tell us about the biology of prokaryotes and eukaryotes, including humans. Methods of DNA sequencing and genome analysis, familiar to many students in a general way, are rapidly improving, and we discuss these advances as well as how bioinformatics is used. This leads to a chapter describing how our knowledge of molecular biology and genetics underpins biotechnology—the application of this knowledge to practical problems and issues such as stem cell research. Part Five closes with a unique sequence of two chapters that explore the interface of developmental processes with molecular biology (Chapter 19) and with evolution (Chapter 20), providing students with a link between these two crucial topics and a bridge to Part Six. PART SIX, THE PATTERNS AND PROCESSES OF EVOLUTION Many students come to the introductory biology course with ideas about evolution already firmly in place. One common view, that evolution is only about Darwin, is firmly put to rest at the start of Chapter 21, which not only illustrates the practical value of fully understanding modern evolutionary biology, but briefly and succinctly traces the history of “Darwin’s dangerous idea” through the twentieth century and up to the present syntheses of molecular evolutionary genetics and evolutionary developmental biology—fields of study that uphold and support the principles of evolutionary biology as the basis for comparing and comprehending all other aspects of biology. The remaining sections of Chapter 21 describe the mechanisms of evolution in clear, matter-of-fact terms. Chapter 22 describes phylogenetic trees as a tool not only of classification but also of evolutionary inquiry. The remaining chapters cover speciation and molecular evolution, concluding with an overview of the evolutionary history of life on Earth. PART SEVEN, THE EVOLUTION OF DIVERSITY Continuing the theme of how evolution has shaped our world, Part Seven introduces the latest views on biodiversity and the evolutionary relationships among organisms. The chapters have been revised with the aim of making it easier for students to appreciate the major evolutionary changes that have taken place within the different groups of organisms. These chapters emphasize understanding the big picture of organismal diversity—the tree of life—as opposed to memorizing a taxonomic hierarchy and names. Throughout the book, the tree of life is emphasized as a way of understanding and organizing biological information. PART EIGHT, FLOWERING PLANTS: FORM AND FUNCTION The emphasis of this modern approach to plant form and function is not only on the basic findings that led to the elucidation of mechanisms for plant growth and reproduction, but also on the use of genetics of model organisms. In response to users of earlier editions, material covering recent discoveries in plant molecular biology and signaling has been reorganized and streamlined to make it more accessible to students. There are also expanded and clearer explanations of such topics as water relations, the plant body plan, and gamete formation and double fertilization. PART NINE, ANIMALS: FORM AND FUNCTION This overview of animal physiology begins with a sequence of chapters covering the systems of information—endocrine, immune, and neural. Learning about these information systems provides important groundwork and explains the processes of control and regulation that affect and integrate the individual physiological systems covered in the remaining chapters of the Part. Chapter 45, “Neurons and Nervous Systems,” has been rearranged and contains descriptions of exciting new discoveries about glial cells and their role in the vertebrate nervous system. The organization of several other chapters has been revised to reflect recent findings and to allow the student to more readily identify the most important concepts to be mastered. PART TEN, ECOLOGY Part Ten continues Life’s commitment to presenting the experimental and quantitative aspects of biology, with increased emphasis on how ecologists design and conduct experiments. New exercises provide opportunities for students to see how ecological data are acquired in the laboratory and in the field, how these data are analyzed, and how the results are applied to answer questions. There is also an expanded discussion of aquatic biomes and a more synthetic explanation of how aquatic, terrestrial, and atmospheric components integrate to influence the distribution and abundance of life on Earth. In addition there is an expanded emphasis on examples of successful strategies proposed by ecologists to mitigate human impacts on the environment; rather than an inventory of ways human activity adversely affects natural systems, this revised Tenth Edition provides more examples of ways that ecological principles can be applied to increase the sustainability of these systems. Exceptional Value Formats We again provide Life both as the full book and as a set of paperback volumes. Thus, instructors who want to use less than the whole book can choose from these split volumes, each of which contains the book’s front matter, appendices, glossary, and index. • Volume I, The Cell and Heredity, includes: Part One, The Science of Life and Its Chemical Basis (Chapters 1–4); Part Two, Cells (Chapters 5–7); Part Three, Cells and Energy (Chapters 8–10); Part Four, Genes and Heredity (Chapters 11–16); and Part Five, Genomes (Chapters 17–20). • Volume II, Evolution, Diversity, and Ecology, includes: Chapter 1, Studying Life; Part Six, The Patterns and Processes of Evolution (Chapters 21–25); Part Seven, The Evolution of Diversity (Chapters 26–33); and Part Ten, Ecology (Chapters 54–59). • Volume III, Plants and Animals, includes: Chapter 1, Studying Life; Part Eight, Flowering Plants: Form and Function (Chapters 34–39); and Part Nine, Animals: Form and Function (Chapters 40–53). Responding to student concerns, there also are two ways to obtain the entire book at a significantly reduced cost. The looseleaf edition of Life is a shrink-wrapped, unbound, three-holepunched version that fits into a three-ring binder. Students take only what they need to class and can easily integrate instructor handouts and other resources. Life was the first comprehensive biology text to offer the entire book as a truly robust eBook, and we offer the Tenth Edition in this flexible, interactive format that gives students a different way to read the text and learn the material. The eBook integrates student media resources (animations, activities, interactive summaries, and quizzes) and offers instructors a powerful way to customize the textbook with their own text, images, web links, and, in BioPortal, quizzes, and other materials. We are proud that our print edition is a greener Life that minimizes environmental impact. Life was the first introductory biology text to be printed on paper earning the Forest Stewardship Council label, the “gold standard in green paper,” and it continues to be manufactured from wood harvested from sustainable forests. Many People to Thank One of the wisest pieces of advice ever given to a textbook author is to “be passionate about your subject, but don’t put your ego on the page.” Considering all the people who looked over our shoulders throughout the process of creating this book, this advice could not be more apt. We are indebted to the many people who help to make this book what it is. First and foremost among these are our colleagues, biologists from over 100 institutions. Before we set pen to paper, we solicited the advice of users of Life’s Ninth Edition, as well as users of other books. These reviewers gave detailed suggestions for improvements. Other colleagues acted as reviewers when the book was almost completed, pointing out inaccuracies or lack of clarity. All of these biologists are listed in the reviewer credits, along with the dozens who reviewed all of the revised assessment resources. Once we began writing, we had the superb advice of a team of experienced, knowledgeable, and patient biologists working as development and line editors. Laura Green of Sinauer Associates headed the team and coordinated her own fine work with that of Jane Murfett, Norma Roche, and Liz Pierson to produce a polished and professional text. We are especially indebted to Laura for her work on the important Investigating Life and new Working with Data elements. For the tenth time in ten editions, Carol Wigg oversaw the editorial process. Her positive influence pervades the entire book. Artist Elizabeth Morales again translated our crude sketches into beautiful new illustrations. We hope you agree that our art program remains superbly clear and elegant. Johannah Walkowicz effectively coordinated the hundreds of reviews described above. David McIntyre, photo editor extraordinaire, researched and provided us with new photographs, including many of his own, to enrich the book’s content and visual statement. Joanne Delphia is responsible for the crisp new design and layout that make this edition of Life not just clear and readable but beautiful as well. Christopher Small headed Sinauer’s production team and contributed in innumerable ways to bringing Life to its final form. Jason Dirks coordinated the creation of our array of media and instructor resources, with Mary Tyler, Mitch Walkowicz, and Carolyn Wetzel serving as editors for our expanded assessment supplements. W. H. Freeman continues to bring Life to a wider audience. Associate Director of Marketing Debbie Clare, the regional specialists, regional managers, and experienced sales force are effective ambassadors and skillful transmitters of the features and unique strengths of our book. We depend on their expertise and energy to keep us in touch with how Life is perceived by its users. Thanks also to the Freeman media group for eBook and BioPortal production. Finally, we thank our friend Andy Sinauer. Like ours, his name is on the cover of the book, and he truly cares deeply about what goes into it. DAVID SADAVA DAVID HILLIS CRAIG HELLER MAY BERENBAUM DAVID SADAVA is the Pritzker Family Foundation Professor of Biology, Emeritus at the Keck Science Center of Claremont McKenna, Pitzer, and Scripps, three of The Claremont Colleges. In addition, he is Adjunct Professor of Cancer Cell Biology at the City of Hope Medical Center in Duarte, California. Twice winner of the Huntoon Award for superior teaching, Dr. Sadava has taught courses on introductory biology, biotechnology, biochemistry, cell biology, molecular biology, plant biology, and cancer biology. In addition to Life: The Science of Biology and Principles of Life, he is the author or coauthor of books on cell biology and on plants, genes, and crop biotechnology. His research has resulted in many papers coauthored with his students, on topics ranging from plant biochemistry to pharmacology of narcotic analgesics to human genetic diseases. For the past 15 years, he has investigated multidrug resistance in human small-cell lung carcinoma cells with a view to understanding and overcoming this clinical challenge. At the City of Hope, his current work focuses on new anti-cancer agents from plants. He is the featured lecturer in “Understanding Genetics: DNA, Genes and their Real-World Applications,“ a video course for The Great Courses series. DAVID M. HILLIS is the Alfred W. Roark Centennial Professor in Integrative Biology and the Director of the Dean’s Scholars Program at the University of Texas at Austin, where he also has directed the School of Biological Sciences and the Center for Computational Biology and Bioinformatics. Dr. Hillis has taught courses in introductory biology, genetics, evolution, systematics, and biodiversity. He has been elected to the National Academy of Sciences and the American Academy of Arts and Sciences, awarded a John D. and Catherine T. MacArthur fellowship, and has served as President of the Society for the Study of Evolution and of the Society of Systematic Biologists. He served on the National Research Council committee that wrote the report BIO 2010: Transforming Undergraduate Biology Education for Research Biologists. His research interests span much of evolutionary biology, including experimental studies of viral evolution, empirical studies of natural molecular evolution, applications of phylogenetics, analyses of biodiversity, and evolutionary modeling. He is particularly interested in teaching and research about the practical applications of evolutionary biology. H. CRAIG HELLER is the Lorry I. Lokey/Business Wire Professor in Biological Sciences and Human Biology at Stanford University. He has taught in the core biology courses at Stanford since 1972 and served as Director of the Program in Human Biology, Chairman of the Biological Sciences Department, and Associate Dean of Research. Dr. Heller is a fellow of the American Association for the Advancement of Science and a recipient of the Walter J. Gores Award for excellence in teaching and the Kenneth Cuthberson Award for Exceptional Service to Stanford University. His research is on the neurobiology of sleep and circadian rhythms, mammalian hibernation, the regulation of body temperature, the physiology of human performance, and the neurobiology of learning. He has done research on a huge variety of animals and physiological problems, including from sleeping kangaroo rats, diving seals, hibernating bears, photoperiodic hamsters, and exercising athletes. Dr. Heller has extended his enthusiasm for promoting active learning via the development of a two-year curriculum in human biology for the middle grades, through the production of Virtual Labs—interactive computer-based modules to teach physiology. MAY BERENBAUM is the Swanlund Professor and Head of the Department of Entomology at the University of Illinois at Urbana- Champaign. She has taught courses in introductory animal biology, entomology, insect ecology, and chemical ecology and has received teaching awards at the regional and national levels from the Entomological Society of America. A fellow of the National Academy of Sciences, the American Academy of Arts and Sciences, and the American Philosophical Society, she served as President of the American Institute for Biological Sciences in 2009 and currently serves on the Board of Directors of AAAS. Her research addresses insect–plant coevolution and ranges from molecular mechanisms of detoxification to impacts of herbivory on community structure. Concerned with the practical application of ecological and evolutionary principles, she has examined impacts of genetic engineering, global climate change, and invasive species on natural and agricultural ecosystems. In recognition of her work, she received the 2011 Tyler Prize for Environmental Achievement. Devoted to fostering science literacy, she has published numerous articles and five books on insects for the general public. The Authors DAVID HILLIS MAY BERENBAUM CRAIG HELLER DAVID SADAVA
Co-published by Sinauer Associates, Inc., and W. H. Freeman and Company. Visit the Life, Eighth Edition preview site. LIFE HAS EVOLVED. . . from its original publication to this dramatically revitalized Eighth Edition. LIFE has always shown students how biology works, offering an engaging and coherent presentation of the fundamentals of biology by describing the landmark experiments that revealed them. This edition builds on those strengths and introduces several innovations. As with previous editions, the Eighth Edition will also be available in three paperback volumes: • Volume I: The Cell and Heredity, Chapters 1-20 • Volume II: Evolution, Diversity and Ecology, Chapters 1, 21-33, 52-57 • Volume III: Plants and Animals, Chapters 1, 34-51
This text aims to establish biology as a discipline, not just a collection of facts. 'Life' develops students' understanding of biological processes with scholarship, a smooth narrative, experimental contexts, art and effective pedagogy.
From its first edition, Life has set the standard for experiment-based introductory biology texts. There is no stronger textbook for helping students understand not just what we know (scientific facts), but how we know it (the experimental process that leads to their discovery). The new edition of Life builds upon this tradition, teaching fundamental concepts and showcasing significant research while responding to changes in biology education... • PEDAGOGICALLY, with features that match the way students learn today, including chapter opening stories, art with balloon captions, and new Learning Objectives • SCIENTIFICALLY, with a wealth of important new research throughout (see Table of Contents for highlights) • TECHNOLOGICALLY, with instant access QR codes printed in the text, new interactive features (media clips, chapter summaries, a flashcard app), and a dramatically enhanced BioPortal, with the adaptive quizzing system, LearningCurve • QUANTIFIABLY, with completely revised assessment resources and new ways of measuring students' progress Also avalable, Volume Splits:—paperbound in full color! Volume 1: The Cell and Heredity (Chapters 1-20) Volume 2: Evolution, Diversity, and Ecology (Chapters 1, 21-33, 54-59) Volume 3: Plants and Animals (Chapters 1, 34-53)
The Eleventh Edition of Life: The Science of Biology is engaging, active, and focused on teaching the skills that students need in the majors biology course. New pedagogical features grab students’ attention and give them a clear learning path through the text. Active learning is a priority throughout the text and media, giving instructors the support they need to encourage students to "learn by doing". Life continues and improves its focus on experiments and data, ensuring that students learn the skills they need to succeed in their careers. It is this potent combination of expertly crafted pedagogy and engagement that make this new edition the best resource for biology students. The Eleventh Edition of Life: The Science of Biology retains its reputation as the book with the highest quality content, clarity of language, and experimental emphasis, and the new focus and features make it a Life worth investigating.

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