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The Fifth Edition of An Introduction to Brain and Behavior continues to reflect the evolution of behavioral neuroscience. In keeping with this evolution, we welcome G. Campbell Teskey, whose fresh perspective—especially on topics related to neurophysiology and nervous system disorders—enhances our author team.

Other major changes in this edition include a deeper emphasis on genetics and epigenetics throughout. Epigenetics is especially important for understanding brain and behavior because environmentally induced modifications in gene expression alter the brain and ultimately behavioral development. Thus, experience—especially early experience—modifies how brain development unfolds. These modifications—of at least some behavioral traits—can be transferred across generations, a process known as epigenetic inheritance. We introduce it in the case study at the end of Section 3-3.

This edition fully addresses advances in imaging technology, including techniques that are fueling the burgeoning field of connectomics and progress toward a comprehensive map of neural connections—a brain connectome. These exciting advances are especially relevant in the second half of the book, where we review higher-level functions.

Imaging advances and epigenetics concepts and research continue as a prime focus in this revision but are not our sole focus. In Chapter 2 we introduce the enteric nervous system, which controls the gut, and later chapters elaborate on ENS functioning. Section 7-1 introduces the emerging field of synthetic neurobiology, elaborated in Section 16-2. Section 5-2 adds new research on lipid neurotransmitters and detail on receptor subtypes.

The range of updates and new coverage in the Fifth Edition text and Focus features is listed, chapter by chapter, in the Preface pages below. See for yourself the breadth and scope of the revision; then read on to learn more about the big-picture improvements in the Fifth Edition.

With encouraging feedback from readers, the book’s learning apparatus continues to feature sets of self-test questions at the end of the major sections in each chapter. These Section Reviews help students track their understanding as they progress. Answers appear at the back of the book.

We continue to expand the popular margin notes. Beyond offering useful asides to the text narrative, these marginalia increase the reader’s ease in finding information, especially when related concepts are introduced early in the text then elaborated on in later chapters. Readers can return quickly to an earlier discussion to refresh their knowledge or jump ahead to learn more. The margin notes also help instructors move through the book to preview later discussions.

The illustrated Experiments, one of the book’s most popular features, show readers how researchers design experiments, that is, how they approach the study of brain–behavior relationships. The Basics features let students brush up or get up to speed on their science foundation—knowledge that helps them comprehend behavioral neuroscience.

We have made some big changes, yet much of the book remains familiar. In shaping content throughout, we continue to examine the nervous system with a focus on function, on how our behavior and our brain interact, by asking key questions that students and neuroscientists ask:

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Every chapter’s central question highlights the brain–behavior relationship. When we first describe how neurons communicate in Section 5-4, for example, we also describe how synaptic plasticity serves as the basis of learning. Later, in Section 14-4, we expand on plasticity as we explore learning and memory.

As it was when we wrote the First Edition, our goal in this new edition is to bring coherence to a vast subject by helping students understand the big picture. Asking fundamental questions about the brain has another benefit: it piques students’ interest and challenges them to join us on the journey of discovery that is brain science.

Scientific understanding of the human brain and human behavior continues to grow at an exponential pace. We want to communicate the excitement of recent breakthroughs in brain science and to relate some of our own experiences from a combined 120+ years of studying brain and behavior, both to make the field’s developing core concepts and latest revelations understandable and to transport uninitiated students to the frontiers of physiological psychology.

To convey the excitement of neuroscience as researchers understand it, we interweave evolution, genetics, and epigenetics; psychopharmacology; and neural plasticity and connectivity, including CNS and ENS interactions, throughout the book.

EVOLUTION Our perspective—neuroscience in an evolutionary context—recurs in almost every chapter. By focusing on comparative behavior and anatomy, we address nervous system evolution in depth in Chapters 1 and 2, evolution of the synapse in Section 5-1, and evolution of visual pathways in Section 9-2. We add ideas about how natural selection might promote overeating in Section 12-5 and evolutionary theories of sleeping and dreaming in Section 13-3. We describe the evolution of sex differences in spatial cognition and language in Section 15-5 and links between our evolved reactions to stress and anxiety disorders in Section 16-4.

GENETICS AND EPIGENETICS We introduce the foundations of genetic and epigenetic research in Sections 1-3 and 2-1 and begin to elaborate on them in Section 3-3. Chapter 5 includes discussions of metabotropic receptors and DNA and of learning and genes. The interplay of genes and drug action is integral to Chapter 6, as are the developmental roles of genes and gene methylation to Chapter 8. Section 9-4 explains the genetics of color vision, and the genetics of sleep disorders anchors Section 13-6. Section 14-4 now includes the role of epigenetics in memory. Sections 16-1 and 16-3 consider the roles of genetics and of prions in understanding the causes of behavioral disorders.

PSYCHOPHARMACOLOGY Chapter 6 investigates how drugs and hormones affect behavior, topics we revisit often through the book. You will find coverage of drugs and information transfer in Section 4-3, drugs and cellular communication in Section 5-3, and synthetic neurobiology in Section 7-1. Section 12-6 covers drugs and motivation; Section 13-6, drugs and sleep disorders; and Section 14-4, neuronal changes with drug use. Section 16-2 discusses the promise of the liposome as a delivery vehicle in pharmacological treatments, and Section 16-4, drugs used as treatments for a range of behavioral disorders.

CONNECTIVITY Neural plasticity is a hallmark of this book. We introduce the concept in Section 1-5, define it in Section 2-1, develop it in Section 2-6, and expand on it throughout. At the conclusion of Section 14-4, we elaborate seven Guiding Principles of Brain Plasticity. We describe the expanding boundaries of connectomics in Section 15-3. The new field of psychotropics, which identifies the connection between the gut microbiome and its effects on the enteric nervous system—as well as on the central nervous system—appears in Sections 2-5 and 12-5.

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We describe the journey of discovery that is neuroscience in a way that students just beginning to study the brain and behavior can understand; then they can use our clinical examples to tie its relevance to the real world. Our approach provides the background students need to understand introductory brain science. Multiple illustrated Experiments in 13 chapters help them visualize the scientific method and how scientists think. The Basics features in 6 chapters address the fact that understanding brain function requires understanding information from all the basic sciences.

These encounters can prove both a surprise and a shock to students who come to the course without the necessary background. The Basics features in Chapters 1 and 2 address the relevant evolutionary and anatomical background. In Chapter 3, The Basics provides a short introduction to chemistry before the text describes the brain’s chemical activities. In Chapter 4 The Basics addresses electricity before exploring the brain’s electrical activity.

Readers already comfortable with the material can easily skip it; less experienced readers can learn it and use it as a context for neuroscience. Students with this background can tackle brain science with greater confidence. Similarly, for students with limited knowledge of basic psychology, we review such facts as stages of behavioral development in Chapter 8 and forms of learning and memory in Chapter 14.

Students in social science disciplines often remark on the amount of biology and chemistry in the book, whereas an equal number of students in biological sciences remark on the amount of psychology. More than half the students enrolled in the Bachelor’s of Science in Neuroscience program at the University of Lethbridge have switched from a biochemistry or psychology major after taking this course. We must be doing something right!

Chapter 7 showcases the range of methods behavioral neuroscientists use to measure and manipulate brain and behavior—traditional methods and such cutting-edge techniques as optical tomography, resting-state fMRI, chemogenetics, and DREADD. Expanded discussions of techniques appear where appropriate, especially in Research Focus features, including Focus 3-2, Brainbow: Rainbow Neurons; Focus 4-3, Optogenetics and Light-Sensitive Channels; and Focus 16-1, Posttraumatic Stress Disorder, which includes treatments based on virtual reality exposure therapies.

Finally, because critical thinking is vital to progress in science, select discussions throughout the book center on relevant aspects. Section 1-2 concludes with The Separate Realms of Science and Belief. Focus 15-3, The Rise and Fall of Mirror Neurons, demonstrates how the media—and even scientists—can fail to question the validity of research results. Section 12-5 introduces the idea that gender identity comprises a broad spectrum rather than a female–male dichotomy. Section 7-7 considers issues of animal welfare in scientific research and the use of laboratory animal models to mimic human neurologic and psychiatric disorders.

Neuroscience is a human science. Everything in this book is relevant to our lives, and everything in our lives is relevant to neuroscience. Understanding neuroscience helps us understand how we learn, how we develop, and how we can help people with brain and behavioral disorders. Knowledge of how we learn, how we develop, and the symptoms of brain and behavioral disorders offer insights into neuroscience.

Clinical material also helps to make neurobiology particularly relevant to students who are going on to a career in psychology, social work, or another profession related to mental health, as well as to students of the biological sciences. We integrate clinical information throughout the text and Clinical Focus features, and we expand on it in Chapter 16, the book’s capstone, as well.

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In An Introduction to Brain and Behavior, the placement of some topics is novel relative to traditional treatments. We include brief descriptions of brain diseases close to discussions of basic associated processes, as exemplified in the integrated coverage of Parkinson disease through Chapter 5, How Do Neurons Communicate and Adapt? This strategy helps first-time students repeatedly forge close links between what they are learning and real-life issues.

To provide a consistent disease nomenclature, the Fifth Edition follows the system advocated by the World Health Organization for diseases named after their putative discoverers. Down syndrome, for example, has largely replaced Down’s syndrome in the popular and scientific literature. We extend that convention to Parkinson disease and Alzheimer disease, among other eponymous diseases and disorders.

This edition covers nearly 150 disorders. Chapter 16 expands on the nature of neuroscience research and the multidisciplinary treatment methods for neurological and psychiatric disorders described in preceding chapters. Its discussion of causes and classifications of abnormal behavior includes updated Table 16-3, now thoroughly revised to conform to the DSM-5 classification.

We emphasize questions that relate to the biological bases of behavior. For us, the excitement of neuroscience lies in understanding how the brain explains what we do, whether it is talking, sleeping, seeing, or learning. Readers will therefore find nearly as many illustrations about behavior as illustrations about the brain. This emphasis on explaining the biological foundation of behavior is another reason that we include a mix of Clinical, Research, and Comparative Focus features throughout the text.

Building on the innovative teaching devices described so far, numerous in-text pedagogical aids adorn every chapter, beginning with an outline and an opening Focus feature that draws students into the chapter’s topic. Focus features dot each chapter to connect brain and behavior to relevant clinical or research experience. Within chapters, definitions of boldface key terms introduced in the text appear in the margins as reinforcement, margin notes link topics together, and end-of-section Review self-tests help students check their grasp of major points.

Each chapter ends with a Summary—several include summarizing tables or illustrations to help students visualize or review big-picture concepts—and a list of Key Terms, each referenced to the page number on which the term is defined. Following this Preface, the Media and Supplements section describes the wide array of supplemental materials designed exclusively for students and teachers using the Fifth Edition.

Our most important learning aid appears on nearly every page in the book: an expansive and, we believe, exceptional set of illustrations. Overwhelmingly, readers agree that, hand in hand with our words, the diagrams describe and illuminate the nervous system. Important anatomical illustrations are large-format to ease perusal. We have retained applications photos that range from a dance class for Parkinson patients in Section 5-3 to a seniors’ bridge game in Section 16-3. New photos include, to illustrate color constancy in Section 9-4, the dress that inspired a social media controversy.

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Illustrations are consistent from chapter to chapter so as to reinforce one another. We consistently color-code diagrams that illustrate each aspect of the neuron, depict each structural region in the brain, and demark nervous system divisions. We include many varieties of micrographic images to show what a particular neural structure actually looks like. These illustrations and images are included on our PowerPoint presentations and integrated as labeling exercises in our Study Guide and Testing materials.

If a textbook is not enjoyable, it has little chance of teaching well. We heighten students’ interest through abundant use of metaphors and examples. Students read about patients whose brain injuries offer insights into brain function, and we examine car engines, robots, and prehistoric flutes for the same purpose. Frequent illustrated Experiments, comparative biology examples, and representative Comparative Focus features help students understand how much we humans have in common with creatures as far distant from us as sea slugs and as close to us as chimpanzees.

We also facilitate learning by reemphasizing main points and by distilling sets of principles about brain function that offer a framework to guide students’ thinking. Thus, Section 2-6 introduces ten key principles that explain how the parts of the nervous system work together. Section 14-4 summarizes seven guiding principles of neuroplasticity. These sets of principles form the basis of many discussions throughout the book. Frequently, marginal notes remind readers when they encounter these principles again—and where to review them in depth.

One challenge in writing an introductory book on any topic is deciding what to include and what to exclude. We organize discussions to focus on the bigger picture—a focus exemplified by the ten principles of nervous system function introduced in Section 2-6 and echoed throughout the book. Any set of principles may be arbitrary yet nevertheless afford students a useful framework for understanding the brain’s activities.

In Chapters 8 through 16 we tackle behavioral topics in a more general way than most contemporary books do. In Chapter 12, for instance, we revisit experiments and ideas from the 1960s to understand why animals behave as they do, then we consider emotional and motivated behaviors as diverse as eating and anxiety attacks in humans. In Chapter 14, the larger picture of learning and memory is presented alongside a discussion of recovery from traumatic brain injury.

This broad focus helps students grasp the big picture that behavioral neuroscience paints. While broadening our focus requires us to leave out some details, our experience with students and teachers through four earlier editions confirms that discussing the larger problems and issues in brain and behavior is of greater interest to students, especially those new to this field, and is more often remembered than are myriad details without context.

As in preceding editions, we are selective in our citation of the truly massive literature on the brain and behavior because we believe that too many citations can disrupt the text’s flow and distract students from the task of mastering concepts. We provide citations to classic works by including the names of the researchers and by mentioning where the research was performed. In areas where controversy or new breakthroughs predominate, we also include detailed citations—177 in all—to papers (especially reviews) from the years 2013 to 2016. An end-of-book References section lists, by chapter, all the literature used in developing the book, reflecting the addition of about 200 new citations total in this edition and elimination of other, now superseded, research.

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As in past editions of this text and Fundamentals of Human Neuropsychology, we have a special debt to Barbara Brooks, our long-time development editor, who has left a strong imprint on both books.

We sincerely thank as well the many people who also contributed to the development of this edition. The staff at Worth Publishers is remarkable and makes revisions a joy to do. We thank our sponsoring editor, Daniel DeBonis, more than ably assisted by Katie Pachnos; our project editor, Edgar Doolan; and production manager Paul Rohloff.

We thank design manager Blake Logan for a striking cover and Charles Yuen for a fresh, inviting, accessible new interior design. Thanks also to Cecilia Varas for coordinating photo research and to Richard Fox, who found photographs and other illustrative materials that we would not have found on our own. We are indebted to Macmillan art manager Matt McAdams, illustration coordinator Janice Donnola, and medical illustrator Evelyn Pence for their excellent work in creating new illustrations.

Our colleagues, too, have helped in the development of every edition. For their contributions in shaping the Fifth Edition, we are especially indebted to the reviewers who provided extensive comments on selected chapters and illustrations: Nancy Blum, California State University, Northridge; Kelly Bordner, Southern Connecticut State University; Benjamin Clark, University of New Mexico; Roslyn Fitch, University of Connecticut; Trevor Gilbert, University of Calgary; Nicholas Grahame, Indiana University–Purdue University Indianapolis; Kenneth Troy Harker, University of New Brunswick; Jason Ivanoff, St. Mary’s University; Dwight Kravitz, The George Washington University; Ralph Lydic, University of Tennessee, Knoxville; Paul Meyer, The State University of New York at Buffalo; Jaime Olavarria, University of Washington; Christopher Robison, Florida State University; Claire Scavuzzo, University of Alberta; Sarah Schock, University of Ottawa; Robert Stackman, Florida Atlantic University; Sandra Trafalis, San Jose State University; Douglas Wallace, Northern Illinois University; Matthew Will, University of Missouri, Columbia; and Harris Philip Zeigler, Hunter College.

Likewise, we continue to be indebted to the colleagues who provided extensive comments on selected chapters and illustrations during the development of the Fourth Edition: Mark Basham, Regis University; Pam Costa, Tacoma Community College; Russ Costa, Westminster College; Renee Countryman, Austin College; Kristen D’Anci, Salem State University; Trevor James Hamilton, Grant MacGewn University; Christian Hart, Texas Woman’s University; Matthew Holahan, Carleton University; Chris Jones, College of the Desert; Joy Kannarkat, Norfolk State University; Jennifer Koontz, Orange Coast College; Kate Makerec, William Paterson University of New Jersey; Daniel Montoya, Fayetteville State University; Barbara Oswald, Miami University of Ohio; Gabriel Radvansky, University of Notre Dame; Jackie Rose, Western Washington University; Steven Schandler, Chapman University; Maharaj Singh, Marquette University; Manda Williamson, University of Nebraska—Lincoln.

We’d also like to thank the reviewers who contributed their thoughts to the Third Edition: Chana Akins, University of Kentucky; Michael Anch, Saint Louis University; Maura Mitrushina, California State University, Northridge; Paul Wellman, Texas A&M University; and Ilsun White, Morehead State University. The methods chapter was new to the Third Edition and posed the additional challenge of taking what easily could read like a seed catalog and making it engaging to readers. We therefore are indebted to Margaret G. Ruddy, The College of New Jersey, and Ann Voorhies, University of Washington, for providing extensive advice on the initial version of Chapter 7.

In addition, we thank the reviewers who provided their thoughts on the Second Edition: Barry Anton, University of Puget Sound; R. Bruce Bolster, University of Winnipeg; James Canfield, University of Washington; Edward Castañeda, University of New Mexico; Darragh P. Devine, University of Florida; Kenneth Green, California State University, Long Beach; Eric Jackson, University of New Mexico; Michael Nelson, University of Missouri, Rolla; Joshua S. Rodefer, University of Iowa; Charlene Wages, Francis Marion University; Doug Wallace, Northern Illinois University; Patricia Wallace, Northern Illinois University; and Edie Woods, Madonna University. Sheri Mizumori, University of Washington, deserves special thanks for reading the entire manuscript for accuracy and providing fresh ideas that proved invaluable.

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Finally, we must thank our tolerant wives for putting up with sudden changes in plans as chapters returned, in manuscript or in proof, with hopes for quick turnarounds. We also thank our colleague Robbin Gibb, who uses the book and has provided much feedback, in addition to our undergraduate and graduate students, technicians, and postdoctoral fellows who kept our research programs moving forward when we were engaged in revising the book.

Bryan Kolb, Ian Q. Whishaw, G. Campbell Teskey