Brain & Behavior: An Introduction to Behavioral Neuroscience Fifth Edition
Flip through this book and you’ll see that its pages are chock-full of facts and applications—just a sampling gleaned from a vast supply that grows too fast for any of us to keep up with and that becomes obsolete just as fast. But sifting through those facts and reporting them is neither the most difficult nor the most important function of a good textbook. A greater challenge is that most students fail to share their instructors’ infatuation with learning; perhaps they lack the genes, or the parental role models, or just the idea that learning can be fun. At any rate, they can find a text like this intimidating, and it is the textbook’s role to change their minds.
The colorful illustrations, case studies, and research vignettes may capture students’ interest, but sparking interest alone is not enough. That’s why we’ve adopted a big-picture approach in writing the text, one that marshals facts into explanations and discards the ones left standing around with nothing to do. When you put facts to work that way, you begin to see students look up and say, “That makes sense,” or “I’ve always wondered about that, but I never thought of it that way,” or “Now I understand what was going on with Uncle Edgar.”
We believe education has the capacity to make a person healthy, happy, and productive, and it makes a society strong. Education realizes that promise when it leads people to inquire and to question, when they learn how to learn. When 45% of the public believes in ghosts, and politics has become a game played by shouting the loudest or telling the most convincing lie, education more than ever needs to teach young people to ask, “Where is the evidence?” and “Is that the only possible interpretation?”
To those who would teach and those who would learn, this book is for you.
To the Student
Brain & Behavior is our attempt to reach out to students, to beckon them into the fascinating world of behavioral neuroscience. These are exceptionally exciting times, comparable in many ways to the renaissance that thrust Europe from the Middle Ages into the modern world. According to the American neurologist Stanley B. Prusiner,
Neuroscience is by far the most exciting branch of science because the brain is the most fascinating object in the universe. Every human brain is different—the brain makes each human unique and defines who he or she is.
We know of no scientific discipline with greater potential to answer the burning questions about ourselves than behavioral neuroscience. We hope this textbook will convey that kind of excitement as you read about discoveries that will revolutionize our understanding of what it means to be human.
We want you to succeed in this course, but, more than that, we want you to learn more than you ever imagined you could and to go away with a new appreciation for the promise of behavioral neuroscience. So, we have a few tips we want to pass along. First, try to sit near the front of the class, because those students usually get the best grades. That is probably because they stay more engaged and are encouraged to ask more questions; but to ask good questions, you should always read the text before you go to class. And so you’ll know where you’re going before you begin to read, take a look at “After reading this chapter, you will be able to,” then skim the chapter subheadings, and read the summary at the end of the chapter. Make sure you can define the terms, and visualize the pathways and brain areas as you come across them. Use the questions in the margins as you go through the text, answer the Concept Check questions, and be sure to test yourself at the end using the Test Your Understanding questions. As you read, pay special attention to the text in blue; these are definitions of the most important terms, which are defined again in the glossary at the end of the book. A “play button” icon like the one you see here will tell you which figures have been animated on the text’s website to help sharpen your understanding. Then, don’t forget to look up some of the books and articles in For Further Reading. Finally, carefully curated lists of web resources at the end of every chapter are fully hyperlinked on the Student Resources site at edge.sagepub.com/garrett5e, directing you to a wealth of additional information on the web. If you are considering a career in behavioral neuroscience, make sure you look at the For Further Thought questions at the end of each chapter. If you do all of these things, you won’t just do better in this course; you will leave saying, “I really got something out of that class!” And when it’s time to take the GRE (or MCAT or VCAT), or talk to your doctor, or interview for a biomedical job, or simply read the Science section of the New York Times, you’ll be using the knowledge you gained in this text.
Stories of Brain & Behavior
We wrote Brain & Behavior with you in mind, so we hope you will let us know where we have done things right and, especially, where we have not. We wish you the satisfaction of discovery and knowledge as you read what we have written for you.
To the Instructor
When Bob wrote the first edition of Brain & Behavior, he had one goal: to entice students into the adventure of behavioral neuroscience. There were other good texts out there, but they read as though they were written for students preparing for further neuroscience courses in graduate school. Those students will find this book adequately challenging, but Bob wrote Brain & Behavior so that anyone interested in behavior, including the newly declared sophomore major or the curious student who has wandered over from the history department, can have the deeper understanding that comes from a biological perspective as they take other courses in psychology.
It is not enough to draw students in with lively writing or by piquing their interest with case studies and telling an occasional story along the way; unless they feel they are learning something significant, they won’t stay—they’ll look for excitement in more traditional places. As Bob wrote, he remembered the text he struggled with in his first behavioral neuroscience class; it wasn’t very interesting because neuroscientists knew much less about the biological underpinnings of behavior than they do now. Since that time, they have learned how the brain changes during learning, they have discovered some of the genes and brain deficiencies that cause schizophrenia, and they are beginning to understand how intricate networks of brain cells produce language, make us intelligent, and help us play the piano or find a mate. In other words, behavioral neuroscience has become a lot more interesting. So the material is there; now it is our job to communicate the excitement we have felt in discovering the secrets of the brain and to make a convincing case that behavioral neuroscience has the power to answer students’ questions about behavior.
A good textbook is all about teaching, but there is no teaching if there is no learning. Over the years, our students have taught us a great deal about what they needed to help them learn. For one thing, we realized how important it is for students to build on their knowledge throughout the course, so we have made several changes from the organization of other texts. First, the chapter on neuronal physiology precedes the chapter on the nervous system, because we believe that you can’t begin to understand the brain until you know how its neurons work. And we reversed the usual order of the vision and audition chapters, because audition provides a friendlier context for introducing the basic principles of sensation and perception. The chapters on addiction, motivation, emotion, and sex follow the introduction to neurophysiology; this was done to build student motivation before tackling sensation and perception. Perhaps more significant, some topics have been moved around among chapters so that they can be developed in a more behaviorally meaningful context. So we discuss language along with audition, the body senses with the mechanisms of movement, the sense of taste in the context of feeding behavior, and olfaction in conjunction with sexual behavior. Most unique, though, is the inclusion of a chapter on the biology of intelligence and another on consciousness. The latter is a full treatment of recent developments in the field, rather than being limited to the usual topics of sleep and split-brain behavior. These two chapters strongly reinforce the theme that behavioral neuroscience is personally relevant and capable of addressing important questions.
Brain & Behavior has several features that will motivate students to learn and encourage them to take an active role in their learning. It engages the student with interest-grabbing opening vignettes; illustrative case studies; and In the News, Application, and Research Spotlight features that take an intriguing step beyond the chapter content. Throughout each chapter, questions in the margins keep the student focused on key points, and a Concept Check at the end of each section serves as a reminder of the important ideas. At the end of each chapter, In Perspective emphasizes the importance and implications of what the student has just read, a summary helps organize that information, and Test Your Understanding questions assess the student’s conceptual understanding as well as factual knowledge. Then, For Further Reading suggestions guide the student in exploring the chapter’s topics more fully, and a list of websites points the way to related information on the Internet. We have found over the years that students who use the study aids in a class are also the best performers in the course.
New in the Fifth Edition
It has been just three years since publication of the fourth edition of Brain & Behavior, but that short time has witnessed unprecedented activity in the neurosciences. We (founding author Bob Garrett and new author Gerald Hough) have reviewed thousands of research articles to ensure that the fifth edition provides students with the most up-to-date coverage of behavioral neuroscience possible. We have undertaken the most ambitious revision of Brain & Behavior since the second edition. We have updated more than 300 references, added 30 new terms, revised more than 50 illustrations, freshened the overall look of the text, and polished its presentation. We completely rewrote several sections, not only to bring them up to date but also to provide better organization and clarity and to shift focus to the most significant aspects. We have, for example, added optogenetics as one of the research techniques in Chapter 4; recast the Chapter 5 discussion of reward, brain plasticity, and learning in addiction; in Chapter 8 developed recent thinking that aggression depends on both testosterone-to-cortisol ratio and the balance of activity among the hypothalamus, amygdala, and prefrontal cortex; revised the discussion of long-term potentiation in Chapter 12 to better emphasize the roles of dopamine, synaptic proteins, and neurogenesis; reconceptualized Chapter 14 to bring the discussion of psychological disorders into line with the fifth edition of the Diagnostic and Statistical Manual of Mental Disorders (DSM-5); and revised the final chapter to reflect current understanding of the sleep and waking circuits. To support these changes, we replaced all the In the News features, updated the Applications, and added a new feature, Research Spotlight, to highlight selected recent developments. In addition, we updated the online material and added new animations to better explain difficult concepts.
In recent years behavioral neuroscience has been dominated by two areas of research, and both continue to gain significance in explaining behavior. The first of these is genetics. New gene associations continue to be discovered, for example, linking autism spectrum disorder to genes for GABA and oxytocin receptors and genes concerned with the formation of synaptic proteins. At the same time, intelligence, schizophrenia, Down syndrome, and Alzheimer’s disease are yielding additional secrets as researchers shift their attention to de novo mutations, copy number variations, and epigenetic influences. Also on the genetic front, the CRISPR technique of gene editing (the American Association for the Advancement of Science’s 2015 breakthrough of the year) is showing great promise for medicine while generating increased concerns about modifying the human gene line.
The second research area, involving neural connectivity and networks, is continuing to reveal how the brain works and, in the case of disorders, how it fails to work. For example, we are beginning to appreciate the role of the salience network in consciousness, by detecting conditions that require attention and switching between the default mode network and the executive network. Other recent studies have found decreased connectivity in the cortex, corpus callosum, and thalamus in bipolar disorder, and both decreased and increased connectivity in different brain areas in depression.
At the same time, we are keenly aware of the need to translate research findings into applications whenever possible. Newly developing treatments featured in the text include the use of anti-inflammatory drugs and memory-inhibiting drugs to fight addiction; and treatment of Alzheimer’s disease with nerve growth factor, brain-growth neurotrophic factor, and the drug aducanumab, which are in phase 1, phase 2, and phase 3 clinical trials, respectively. We also applaud the good news in two recent reports that prevalence of autism spectrum disorder appears to have leveled off and the risk for Alzheimer’s disease has declined in several countries, including the United Kingdom (by 20%) and the United States (by 26%).
Revising a textbook like Brain & Behavior is incredibly hard work, and in spite of Bob’s and Gerald’s best efforts, the fifth edition would not be possible without the help of many others. Kudos to acquisitions editor Abbie Rickard and content development editor Lucy Berbeo for their insights, their patience, and their ability to keep us on schedule (almost). Aside from their challenges in dealing with the authors, they have managed a host of contributors, including Meghan C. Kahn, of Indiana University Southeast, who revised the Study Guide and wrote the In the News features; Joshua S. Rodefer, of Valdosta State University, who contributed Application and Research Spotlight features; Richard J. Addante, of California State University, San Bernardino, who reviewed the artwork; and an untold number of illustrators and animators. We also want to thank others who have made the project go so smoothly, including production editors David Felts and Kelly DeRosa, editorial assistant Jennifer Cline, cover designer Gail Buschman, marketing manager Katherine Hepburn, and marketing associate Tzveta Mihaylov.
Bob has had a number of mentors along the way, to whom he is forever grateful. A few of those special people are Wayne Kilgore, who taught the joys of science along with high school chemistry and physics; Garvin McCain, who introduced him to the satisfactions of research; Roger Kirk, who taught him that anything worth doing is worth doing over and over until it’s right; and Ellen Roye and Ouilda Piner, who shared their love of language. These dedicated teachers showed him that learning was his responsibility, and they shaped his life with their unique gifts and quiet enthusiasm.
But of all of Bob’s supporters, the most important was his wife, Duejean; love and thanks to her for fond memories and for her patient understanding and her appreciation of how important this project has been to him.
For Gerald, becoming a coauthor on this textbook, which he has used in his classes over the past decade, is an amazing privilege. He hopes that the instructors and students who use this book find that the overall presentation and helpfulness in assisting in the learning process are unchanged from the previous editions. His background in behavioral neuroethology (a mouthful for sure!) provides a background in understanding the neural bases of behavior. He would like to thank his academic mentors who have helped him find his way in a complex scientific world: Erich Klinghammer at Wolf Park in Battle Ground, Indiana, sowed the seeds for a love of wolves and understanding behavior in naturalistic settings; Susan Volman at The Ohio State University introduced him to the complexities of behavioral neuroscience through their investigation of how birds learn (and forget) songs at the neural level; and Verner Bingman gave him the opportunity to develop as a scholar and mentor to his students at Bowling Green State University in Ohio. He would also like to thank his family for letting him work on this important textbook: his wife of 25 years, Kerry; sons Alexander and Benjamin; and of course Melbourne the parakeet.
In addition, the following reviewers gave generously of their time and expertise throughout the development of this text; they contributed immensely to the quality of Brain & Behavior:
First edition: Susan Anderson, University of South Alabama; Patrizia Curran, University of Massachusetts–Dartmouth; Lloyd Dawe, Cameron University; Tami Eggleston, McKendree College; James Hunsicker, Southwestern Oklahoma State University; Eric Laws, Longwood College; Margaret Letterman, Eastern Connecticut State University; Doug Matthews, University of Memphis; Grant McLaren, Edinboro University of Pennsylvania; Rob Mowrer, Angelo State University; Anna Napoli, University of Redlands; Robert Patterson, Washington State University; Joseph Porter, Virginia Commonwealth University; Jeffrey Stern, University of Michigan–Dearborn; Aurora Torres, University of Alabama in Huntsville; Michael Woodruff, East Tennessee State University; and Phil Zeigler, Hunter College.
Second edition: M. Todd Allen, University of Northern Colorado; Patricia A. Bach, Illinois Institute of Technology; Wayne Brake, University of California–Santa Barbara; Steven I. Dworkin, University of North Carolina; Sean Laraway, San Jose State University; Mindy J. Miserendino, Sacred Heart University; Brady Phelps, South Dakota State University; Susan A. Todd, Bridgewater State College; and Elizabeth Walter, University of Oregon.
Third edition: John A. Agnew, University of Colorado Boulder; Michael A. Bock, American International College; Rachel E. Bowman, Sacred Heart University; Jessica Cail, Pepperdine University; Mary Jo Carnot, Chadron State College; Cheryl A. Frye, The University at Albany–State University of New York; Rebecca L. M. Fuller, Catholic University of America; Cindy Gibson, Washington College; Bennet Givens, Department of Psychology, The Ohio State University; Robert B. Glassman, Lake Forest College; Gerald E. Hough, Rowan University; Joseph Nuñez, Michigan State University; and Kimberly L. Thomas, University of Central Oklahoma.
Fourth edition: John A. Agnew, University of Colorado Boulder; Ben Allen, University of Pittsburgh; Scott L. Decker, University of South Carolina; Carol L. DeVolder, St. Ambrose University; Jeff Dyche, James Madison University; Cindy Gibson, Washington College; Deirdre C. Greer, Columbus State University; William Meil, Indiana University of Pennsylvania; Samar Saade Needham, California State University, Long Beach; M. Foster Olive, Arizona State University; Catherine Powers Ozyurt, Bay Path College; Allen Salo, University of Maine at Presque Isle; Justin P. Smith, University of South Dakota; Gretchen Sprow, University of North Carolina at Chapel Hill; and Sandra Trafalis, San Jose State University.
Fifth edition: Kurt T. Choate, Northeastern State University; Andrea L. O. Hebb, Saint Mary’s University; Bryant Horowitz, East Los Angeles College, California State University, Northridge, and The Chicago School of Professional Psychology; Dwight J. Kravitz, The George Washington University; and Lorenz S. Neuwirth, SUNY College at Old Westbury.
—Bob Garrett and Gerald Hough
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