FULL CONTENTS Major Memory Systems of the Mammalian Brain 1156 Effects of damage to the prefrontal cortex in Behavior Supported by Multiple Memory Systems 1174 Monkeys 1208 lon Neurophysiology of the Prefrontal Cortex 1211 References 1175 Theories of Prefrontal Cortex Function 1217 Suggested Readings 1177 Further readings 1221 51. Language and Communication Suggested Readings 122 DAVID N. CAPLAN AND JAMES L. GOULD Animal Communication 1179 53. Consciousness Human Language 1184 CHRISTOF KOCH Conclusions 1196 References 1197 What Phenomena Does Consciousness Encompass? 1224 Suggested Readings 1198 The Neurobiology of Free Will 1224 Consciousness in Other Species 1225 52. The Prefrontal Cortex and Executive Brain Arousal and States of Consciousness 1225 Fu Inctions The Neuronal Correlates of Consciousness 1228 EARL MILLER AND JONATHAN WALLIS The Neuronal Basis of Perceptual Illusions 1229 Other Perceptual Puzzles of Contemporary Interest 1231 Introduction 1199 Forward versus Feedback Projections 1232 Controlled Processing 1199 An Information-Theoretical Theory of Anatomy and Organization of the Prefrontal Consciousness 1233 Cortex 1201 Conclusion 1234 Effects of Damage to the Prefrontal Cortex in References 1234 Humans 1203 Neuroimaging Studies and PFC 1207 Index 1237
xiv FULL CONTENTS Major Memory Systems of the Mammalian Brain 1156 Behavior Supported by Multiple Memory Systems 1174 Conclusion 1175 References 1175 Suggested Readings 1177 51. Language and Communication DAVID N. CAPLAN AND JAMES L. GOULD Animal Communication 1179 Human Language 1184 Conclusions 1196 References 1197 Suggested Readings 1198 52. The Prefrontal Cortex and Executive Brain Functions EARL MILLER AND JONATHAN WALLIS Introduction 1199 Controlled Processing 1199 Anatomy and Organization of the Prefrontal Cortex 1201 Effects of Damage to the Prefrontal Cortex in Humans 1203 Neuroimaging Studies and PFC 1207 Effects of Damage to the Prefrontal Cortex in Monkeys 1208 Neurophysiology of the Prefrontal Cortex 1211 Theories of Prefrontal Cortex Function 1217 Further Readings 1221 References 1221 Suggested Readings 1221 53. Consciousness CHRISTOF KOCH What Phenomena Does Consciousness Encompass? 1224 The Neurobiology of Free Will 1224 Consciousness in Other Species 1225 Arousal and States of Consciousness 1225 The Neuronal Correlates of Consciousness 1228 The Neuronal Basis of Perceptual Illusions 1229 Other Perceptual Puzzles of Contemporary Interest 1231 Forward versus Feedback Projections 1232 An Information-Theoretical Theory of Consciousness 1233 Conclusion 1234 References 1234 Index 1237
Preface to the Third Edition In this third edition of Fundamental Neuroscience, nervous systems. The remainder of the volume(Sec- ve have tried to improve on the second edition with tions Il-VID) presents the major topics of neuroscience a volume that effectively introduces students to the The second section( Cellular and Molecular Neurosci- full range of contemporary neuroscience. Neuro- ence)considers the cellular and subcellular organiza cience is a large field founded on the premise that all tion of neurons, the physiology of nerve cells, and how of behavior and all of mental life have their origin in signaling occurs between neurons. The third section the structure and function of the nervous system.(Nervous System Development)includes discussion of Today, the need for a single-volume introduction to neural induction, cell fate, migration, process out neuroscience is greater than ever. Towards the end of growth, development of dendrites, synapse formation, the 20 century, the study of the brain moved from a programmed cell death, synapse elimination, and early peripheral position within both the biological and experience including critical periods. The fourth and psychological sciences to become an interdisciplinary fifth sections(Sensory Systems and Motor Systems field that is now central within each discipline. The describe the neural organization of each sensory maturation of neuroscience has meant that individu- modality and the organization of the brain pathway als from diverse backgrounds-including molecular and systems important for locomotion, voluntary biologists, computer scientists, and psychologists-are action, and eye movements. The sixth section(Regula interested in learning about the structure and func- tory Systems) describes the variety of hypothalamic tion of the brain and about how the brain works In and extra-hypothalamic systems that support motiva- addition, new techniques and tools have become tion, reward, and internal regulation, including cardio- available to study the brain in increasing detail. In the vascular function, respiration, food and water intake last 15 years new genetic methods have been intro neuroendocrine function, circadian rhythms, and duced to delete or over-express single genes with and dreaming. The final section( Behavioral and Cog- spatial and temporal specificity. Neuroimaging nitive Neuroscience) describes the neural foundations techniques such as functional magnetic resonance of the so-called higher mental functions including per imaging(fMRI have been developed that allow study ception, attention, memory, language, spatial cogni of the living human brain while it is engaged in tion, and executive function. Additional chapters cover human brain evolution, cognitive development and This third edition attempts to capture the promise aging and consciousness the volume will be accom- and excitement of this fast-moving discipline. All the panied by an easily accessible companion website, chapters have been rewritten to make them more which will present all the figures and increase the flex oncise. As a result the new edition is about 30% ibility with which the material can be used. shorter than previous editions but still covers the same The authors listed at the ends of the chap comprehensive range of topics. The volume begins and boxes are working scientists, experts in the with an opening chapter that provides an overview of topics they cover. The Editors edited the chapters to the discipline. A second chapter presents fundamental achieve consistency of style and content. At Academic information about the architecture and anatomy of Press/Elsevier Science, the project was coordinated
Preface to the Third Edition xv In this third edition of Fundamental Neuroscience, we have tried to improve on the second edition with a volume that effectively introduces students to the full range of contemporary neuroscience. Neuroscience is a large fi eld founded on the premise that all of behavior and all of mental life have their origin in the structure and function of the nervous system. Today, the need for a single-volume introduction to neuroscience is greater than ever. Towards the end of the 20th century, the study of the brain moved from a peripheral position within both the biological and psychological sciences to become an interdisciplinary fi eld that is now central within each discipline. The maturation of neuroscience has meant that individuals from diverse backgrounds—including molecular biologists, computer scientists, and psychologists—are interested in learning about the structure and function of the brain and about how the brain works. In addition, new techniques and tools have become available to study the brain in increasing detail. In the last 15 years new genetic methods have been introduced to delete or over-express single genes with spatial and temporal specifi city. Neuroimaging techniques such as functional magnetic resonance imaging (fMRI) have been developed that allow study of the living human brain while it is engaged in cognition. This third edition attempts to capture the promise and excitement of this fast-moving discipline. All the chapters have been rewritten to make them more concise. As a result the new edition is about 30% shorter than previous editions but still covers the same comprehensive range of topics. The volume begins with an opening chapter that provides an overview of the discipline. A second chapter presents fundamental information about the architecture and anatomy of nervous systems. The remainder of the volume (Sections II–VII) presents the major topics of neuroscience. The second section (Cellular and Molecular Neuroscience) considers the cellular and subcellular organization of neurons, the physiology of nerve cells, and how signaling occurs between neurons. The third section (Nervous System Development) includes discussion of neural induction, cell fate, migration, process outgrowth, development of dendrites, synapse formation, programmed cell death, synapse elimination, and early experience including critical periods. The fourth and fi fth sections (Sensory Systems and Motor Systems) describe the neural organization of each sensory modality and the organization of the brain pathways and systems important for locomotion, voluntary action, and eye movements. The sixth section (Regulatory Systems) describes the variety of hypothalamic and extra-hypothalamic systems that support motivation, reward, and internal regulation, including cardiovascular function, respiration, food and water intake, neuroendocrine function, circadian rhythms, and sleep and dreaming. The fi nal section (Behavioral and Cognitive Neuroscience) describes the neural foundations of the so-called higher mental functions including perception, attention, memory, language, spatial cognition, and executive function. Additional chapters cover human brain evolution, cognitive development and aging, and consciousness. The volume will be accompanied by an easily accessible companion website, which will present all the fi gures and increase the fl exibility with which the material can be used. The authors listed at the ends of the chapters and boxes are working scientists, experts in the topics they cover. The Editors edited the chapters to achieve consistency of style and content. At Academic Press/Elsevier Science, the project was coordinated
PREFACE TO THE THIRD EDITION by Hilary Rowe and Nikki Levy(Publishing Editors), The Editors of Fundamental Neuroscience hope that and we are grateful to them for their leadership and users of this book, and especially the students who will advice throughout the project. In addition, Meg Day become the next generation of neuroscientists, find (Developmental Editor) very capably coordinated the subject matter of neuroscience as interesting and excit- production of the book with the help of Sarah Hajduk ing as we do (Publishing Services Manager)and Christie Jozwiak (Project Manager) The editors
by Hilary Rowe and Nikki Levy (Publishing Editors), and we are grateful to them for their leadership and advice throughout the project. In addition, Meg Day (Developmental Editor) very capably coordinated the production of the book with the help of Sarah Hajduk (Publishing Services Manager) and Christie Jozwiak (Project Manager). xvi PREFACE TO THE THIRD EDITION The Editors of Fundamental Neuroscience hope that users of this book, and especially the students who will become the next generation of neuroscientists, fi nd the subject matter of neuroscience as interesting and exciting as we do. The Editors
About the editors Larry R Squire is Distinguished Professor of Psy- of Systems Neurobiology at the Salk Institute for Bio. chiatry, Neurosciences, and Psychology at the Univer- logical Studies. Her research interests are in the neu- ity of California School of Medicine, San Diego, and robiology of resilience and learning, and her laboratory Research Career Scientist at the Veterans Affairs investigates behavioral, circuit, cellular, and molecular ledical Center, San Diego. He investigates the organi- mechanisms in the sense of balance zation and neurological foundations of memory. He is Anirvan Ghosh is Stephen Kuffler Professor in the a former President of the Society for Neuroscience and Division of Biological Sciences at the University of is a member of the National Academy of Sciences and California, San Diego and Director of the graduate he Institute of medicine program in Neurosciences. His research interests parwin K. Berg is Distinguished Professor in the include the development of synaptic connections in Division of Biological Sciences at the University of the central nervous system and the role of activity California, San Diego. He has been chairman of the dependent gene expression in the cortical develop- Biology Department and currently serves as Councilor ment. He is recipient of the Presidential Early Career of the Society for Neuroscience and as a Board member Award for Scientists and Engineers and the Society for of the Kavli Institute for Brain and Mind. His research Neuroscience Young Investigator Award is focused on the roles of nicotinic cholinergic signal Nicholas C. Spitzer is Distinguished Professor in ing in the vertebrate nervous system. the Division of Biological Sciences at the University Floyd Bloom is Professor Emeritus in the Molecular of California, San Diego. His research is focused and Integrative Neuroscience Department(MIND)at on neuronal differentiation and the role of electrical The Scripps Research Institute. His recent awards activity and calcium signaling in the assembly of include the Sarnat Award from the Institute of Medi- the nervous system. He has been chairman of cine and the Salmon Medal of the New York Academy the Biology Department and the Neurobiology Section, of Medicine. He is a former President of the Society for a trustee of the Grass Foundation, and served as Academy of Sciences and the Institute of Medicine al Councilor of the Society for Neuroscience. He is a Neuroscience and is a member of the nati member of the American Academy of Arts and Sci- Sascha du Lac is an Investigator of the Howard ences and Co-Director of the Kavli Institute for Brain Hughes Medical Institute and an Associate Professor and mind
About The Editors xvii Larry R. Squire is Distinguished Professor of Psychiatry, Neurosciences, and Psychology at the University of California School of Medicine, San Diego, and Research Career Scientist at the Veterans Affairs Medical Center, San Diego. He investigates the organization and neurological foundations of memory. He is a former President of the Society for Neuroscience and is a member of the National Academy of Sciences and the Institute of Medicine. Darwin K. Berg is Distinguished Professor in the Division of Biological Sciences at the University of California, San Diego. He has been chairman of the Biology Department and currently serves as Councilor of the Society for Neuroscience and as a Board member of the Kavli Institute for Brain and Mind. His research is focused on the roles of nicotinic cholinergic signaling in the vertebrate nervous system. Floyd Bloom is Professor Emeritus in the Molecular and Integrative Neuroscience Department (MIND) at The Scripps Research Institute. His recent awards include the Sarnat Award from the Institute of Medicine and the Salmon Medal of the New York Academy of Medicine. He is a former President of the Society for Neuroscience and is a member of the National Academy of Sciences and the Institute of Medicine. Sascha du Lac is an Investigator of the Howard Hughes Medical Institute and an Associate Professor of Systems Neurobiology at the Salk Institute for Biological Studies. Her research interests are in the neurobiology of resilience and learning, and her laboratory investigates behavioral, circuit, cellular, and molecular mechanisms in the sense of balance. Anirvan Ghosh is Stephen Kuffl er Professor in the Division of Biological Sciences at the University of California, San Diego and Director of the graduate program in Neurosciences. His research interests include the development of synaptic connections in the central nervous system and the role of activitydependent gene expression in the cortical development. He is recipient of the Presidential Early Career Award for Scientists and Engineers and the Society for Neuroscience Young Investigator Award. Nicholas C. Spitzer is Distinguished Professor in the Division of Biological Sciences at the University of California, San Diego. His research is focused on neuronal differentiation and the role of electrical activity and calcium signaling in the assembly of the nervous system. He has been chairman of the Biology Department and the Neurobiology Section, a trustee of the Grass Foundation, and served as Councilor of the Society for Neuroscience. He is a member of the American Academy of Arts and Sciences and Co-Director of the Kavli Institute for Brain and Mind
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