Fundamental Neuroscience Third Edition Larry R Squire. Floyd E Bloom. Nicholas C Spitzer Sascha du Lac Anirvan Ghosh.Darwin Berg
Short Contents 18. Target Selection, Topographic Maps, and Synapse Formation 401 NEUROSCIENCE 19. Programmed Cell Death and Neurotrophic 1. Fundamentals of Neuroscience 3 2. Basic Plan of the Nervous System 15 20. Synapse Elimination 469 21. Dendritic Development 491 22. Early Experience and Sensitive Periods 517 CELLULAR AND MOLECULAR NEUROSCIENCE SENSORY SYSTEMS 3. Cellular Components of Nervous Tissue 41 23. Fundamentals of Sensory Systems 535 4. Subcellular Organization of the Nervous System 24. Chemical Senses: Taste and Olfaction 549 25. Somatosensory System 581 5. Electrotonic Properties of Axons and Dendrites 87 26. Audition 609 6. Membrane potential and Action potential 111 27. Vision 637 7. Neurotransmitters 133 8. Release of Neurotransmitters 157 9. Neurotransmitter Receptors 181 10. Intracellular Signaling 205 1. Postsynaptic Potentials and Synaptic MOTOR SYSTEMS Integration 227 12. Complex Information Processing in Dendrites 247 28. Fundamentals of Motor Systems 663 13. Brain Energy Metabolism 271 29. The Spinal and Peripheral Motor System 677 30. Descending Control of Movement 699 31. The Basal Ganglia 725 33. Eye Movements 775 NERVOUS SYSTEM DEVELOPMENT 14. Neural Induction and Pattern Formation 297 REGULATORY SYSTEMS 15. Cellular Determination 321 16. Neurogenesis and Migration 351 34. The Hypothalamus: An Overview of Regulatory 17. Growth Cones and Axon Pathfind
Short Contents v I NEUROSCIENCE 1. Fundamentals of Neuroscience 3 2. Basic Plan of the Nervous System 15 II CELLULAR AND MOLECULAR NEUROSCIENCE 3. Cellular Components of Nervous Tissue 41 4. Subcellular Organization of the Nervous System: Organelles and Their Functions 59 5. Electrotonic Properties of Axons and Dendrites 87 6. Membrane Potential and Action Potential 111 7. Neurotransmitters 133 8. Release of Neurotransmitters 157 9. Neurotransmitter Receptors 181 10. Intracellular Signaling 205 11. Postsynaptic Potentials and Synaptic Integration 227 12. Complex Information Processing in Dendrites 247 13. Brain Energy Metabolism 271 III NERVOUS SYSTEM DEVELOPMENT 14. Neural Induction and Pattern Formation 297 15. Cellular Determination 321 16. Neurogenesis and Migration 351 17. Growth Cones and Axon Pathfi nding 377 18. Target Selection, Topographic Maps, and Synapse Formation 401 19. Programmed Cell Death and Neurotrophic Factors 437 20. Synapse Elimination 469 21. Dendritic Development 491 22. Early Experience and Sensitive Periods 517 IV SENSORY SYSTEMS 23. Fundamentals of Sensory Systems 535 24. Chemical Senses: Taste and Olfaction 549 25. Somatosensory System 581 26. Audition 609 27. Vision 637 V MOTOR SYSTEMS 28. Fundamentals of Motor Systems 663 29. The Spinal and Peripheral Motor System 677 30. Descending Control of Movement 699 31. The Basal Ganglia 725 32. Cerebellum 751 33. Eye Movements 775 VI REGULATORY SYSTEMS 34. The Hypothalamus: An Overview of Regulatory Systems 795
SHORT CONTENTS 35. Central Control of Autonomic Functions: VII Organization of the autonomic Nervous System 807 BEHAVIORAL AND COGNITIVE 36. Neural Regulation of the Cardiovascular NEUROSCIENCE 37. Neural Control of Breathing 855 44. Human Brain Evolution 1019 38. Food Intake and Metabolism 873 45. Cognitive Development and Aging 1039 39. Water Intake and Body Fluids 889 46. Visual Perception of Objects 1067 40. Neuroendocrine Systems 905 47. Spatial 41. Circadian Timekeeping 931 48. Attention 1113 2. Sleep, Dreaming, and Wakefulness 959 49. Learning and Memory: Basic Mechanisms 1133 43. Reward, Motivation, and Addiction 987 0. Learning and Memory: Brain Systems 1153 51. Language and Communication 1179 52. The prefrontal Cortex and Executive brain Functions 1199 53. The Neuroscience of consciousness 1223
35. Central Control of Autonomic Functions: Organization of the Autonomic Nervous System 807 36. Neural Regulation of the Cardiovascular System 829 37. Neural Control of Breathing 855 38. Food Intake and Metabolism 873 39. Water Intake and Body Fluids 889 40. Neuroendocrine Systems 905 41. Circadian Timekeeping 931 42. Sleep, Dreaming, and Wakefulness 959 43. Reward, Motivation, and Addiction 987 VII BEHAVIORAL AND COGNITIVE NEUROSCIENCE 44. Human Brain Evolution 1019 45. Cognitive Development and Aging 1039 46. Visual Perception of Objects 1067 47. Spatial Cognition 1091 48. Attention 1113 49. Learning and Memory: Basic Mechanisms 1133 50. Learning and Memory: Brain Systems 1153 51. Language and Communication 1179 52. The Prefrontal Cortex and Executive Brain Functions 1199 53. The Neuroscience of Consciousness 1223 vi SHORT CONTENTS
Full contents Preface xv Development Reveals Basic Vertebrate Parts 22 About the editors xvii The Basic Plan of Nervous System Connectivity 27 List of contributors xix Overview of the adult mammalian nervous System 31 References 37 Suggested Readings 38 NEUROSCIENCE 1. Fundamentals of neuroscience CELLULAR AND MOLECULAR FLOYD E BLOOM NEUROSCIENCE A Brief History of Neuroscience 3 The Terminology of Nervous Systems Is Hierarchical, 3. Cellular Components of Distributed, Descriptive, and Historically Based 3 Nervous Tissue Neurons and glia are cellular building blocks of the PATRICK R. HOF, JEAN DE VELLIS, ESTHER A NIMCHINSKY Nervous System 4 GRAHAME KIDD, LUZ CLAUDIO, AND BRUCE D. TRAPP The Operative Processes of Nervous Systems Are als Hierarchical 5 Neurons 41 Cellular Organization of the brain 6 Specific Examples of Different Neuronal Types 45 Organization of this Text 7 Neuroglia 47 This Book Is Intended for a Broad Range of Scholars of Cerebral Vasculature 54 the neurosciences 8 References 57 Clinical Issues in the neurosciences 8 Suggested Readings 58 The Spirit of Exploration Con Ines The Genomic Inventory Is a Giant Step Forward 9 4. Subcellular Organization of the Neuroscience Today: A Communal Endeavor 10 ervous system s an The Creation of Knowledge 10 Their functions Responsible Conduct 11 SCOTT T BRADY, DAVIDR. COLMAN, AND PETER J. BROPHY Summary 13 References 13 Axons and Dendrites: Unique Structural Components of eurons 2. Basic Plan of the Nervous System Protein Synthesis in Nervous Tissue 63 LARRY W. SWANSON Cytoskeletons of Neurons and Glial Cells 70 Molecular Motors in the Nervous System 77 Introduction 15 Building and Maintaining Nervous System Cells 80 Evolution Highlights: General Organizing Principles 15 Refe rences
Full Contents vii Preface xv About the Editors xvii List of Contributors xix I NEUROSCIENCE 1. Fundamentals of Neuroscience FLOYD E. BLOOM A Brief History of Neuroscience 3 The Terminology of Nervous Systems Is Hierarchical, Distributed, Descriptive, and Historically Based 3 Neurons and Glia Are Cellular Building Blocks of the Nervous System 4 The Operative Processes of Nervous Systems Are also Hierarchical 5 Cellular Organization of the Brain 6 Organization of this Text 7 This Book Is Intended for a Broad Range of Scholars of the Neurosciences 8 Clinical Issues in the Neurosciences 8 The Spirit of Exploration Continues 9 The Genomic Inventory Is a Giant Step Forward 9 Neuroscience Today: A Communal Endeavor 10 The Creation of Knowledge 10 Responsible Conduct 11 Summary 13 References 13 2. Basic Plan of the Nervous System LARRY W. SWANSON Introduction 15 Evolution Highlights: General Organizing Principles 15 Development Reveals Basic Vertebrate Parts 22 The Basic Plan of Nervous System Connectivity 27 Overview of the Adult Mammalian Nervous System 31 References 37 Suggested Readings 38 II CELLULAR AND MOLECULAR NEUROSCIENCE 3. Cellular Components of Nervous Tissue PATRICK R. HOF, JEAN DE VELLIS, ESTHER A. NIMCHINSKY, GRAHAME KIDD, LUZ CLAUDIO, AND BRUCE D. TRAPP Neurons 41 Specifi c Examples of Different Neuronal Types 45 Neuroglia 47 Cerebral Vasculature 54 References 57 Suggested Readings 58 4. Subcellular Organization of the Nervous System: Organelles and Their Functions SCOTT T. BRADY, DAVID R. COLMAN, AND PETER J. BROPHY Axons and Dendrites: Unique Structural Components of Neurons 59 Protein Synthesis in Nervous Tissue 63 Cytoskeletons of Neurons and Glial Cells 70 Molecular Motors in the Nervous System 77 Building and Maintaining Nervous System Cells 80 References 85
ⅶi FULL CONTENTS 5. Electrotonic Properties of Axons and 9. Neurotransmitter Receptors Dendrites M. NEAL WAXHAM GORDON M SHEPHERD ionotropic Receptors 181 Toward a Theory of Neuronal Information G-Protein Coupled Receptors 193 Processing 87 References 203 Basic Tools: Cable Theory and Compartmental Models 88 10. Intracellular Signaling Spread of Steady-State Signals 88 HOWARD SCHULMAN AND JAMES L. ROBERTS Spread of Transient Signals 93 Electrotonic Properties Underlying Propagation in Signaling Through G-Protein-Linked Receptors 205 Axons 95 Modulation of Neuronal Function by Protein Kinases and Electrotonic Spread in Dendrites 98 Phosphatases 214 Dynamic Properties of Passive Electrotonic Intracellular Signaling Affects Nuclear Gene Structure 101 Expression 222 Relating Passive to Active Potentials 106 References 226 Suggested Readings 226 6. Membrane potential and 11. Postsynaptic Potentials and Synaptic Action po DAVID A MCCORMICK JOHN H. BYRNE Membrane potential 112 ionotropic Receptors: Mediators of Fast Excitatory and Action Potential 117 Inhibitory S Synaptic Potentials 227 erences Metabotropic Receptors: Mediators of Slow Synapt Suggested Readings 132 Potentials 239 Integration of Synaptic Potentials 242 References 245 7. Neurotransmitters Suggested Readings 245 ARIEL Y DEUTCH AND ROBERT H. ROTH 12. Complex Information Processing in Several modes of neuronal communication Dendrites Exist 133 Chemical Transmission 134 GORDXON M. SHEPHERD Classical Neurotransmitters 136 Nonclassical Neurotransmitters 147 Strategies for Studying Complex Dendrites 247 Peptide Transmitters 148 Building Principles Step by Step 248 An Axon places Constraints on dendritic conven Itional Transmitters 149 Synaptic Transmission in Perspective 154 Processing 249 References 154 Dendrodendritic Interactions between Axonal Cells 250 Passive Dendritic Trees Can Perform Complex Computations 251 8. Release of neurotransmitters Separation of Dendritic Fields Enhances Complex THOMAS L SCHWARZ Information Processing 252 Distal Dendrites Can be Closely Linked to Axonal Transmitter Release Is Quantal 157 Output 253 Excitation-Secretion Coupling 160 Depolarizing and Hyperpolarizing Dendritic Conductances Molecular mechanisms of the nerve Interact Dynamically 255 The Axon Hillock-Initial Segment Encodes Global Quantal Analysis: Probing Synaptic Output 256 hysiology 173 Short-Term Synaptic Plasticity 176 References 180 e Controy talse Initiation Sites Are under Dynamic Multiple lmi etrograde Impulse Spread into Dendrites Can have Suggested Readings 180
viii FULL CONTENTS 5. Electrotonic Properties of Axons and Dendrites GORDON M. SHEPHERD Toward a Theory of Neuronal Information Processing 87 Basic Tools: Cable Theory and Compartmental Models 88 Spread of Steady-State Signals 88 Spread of Transient Signals 93 Electrotonic Properties Underlying Propagation in Axons 95 Electrotonic Spread in Dendrites 98 Dynamic Properties of Passive Electrotonic Structure 101 Relating Passive to Active Potentials 106 References 108 6. Membrane Potential and Action Potential DAVID A. MCCORMICK Membrane Potential 112 Action Potential 117 References 131 Suggested Readings 132 7. Neurotransmitters ARIEL Y. DEUTCH AND ROBERT H. ROTH Several Modes of Neuronal Communication Exist 133 Chemical Transmission 134 Classical Neurotransmitters 136 Nonclassical Neurotransmitters 147 Peptide Transmitters 148 Unconventional Transmitters 149 Synaptic Transmission in Perspective 154 References 154 8. Release of Neurotransmitters THOMAS L. SCHWARZ Transmitter Release Is Quantal 157 Excitation–Secretion Coupling 160 Molecular Mechanisms of the Nerve Terminal 163 Quantal Analysis: Probing Synaptic Physiology 173 Short-Term Synaptic Plasticity 176 References 180 Suggested Readings 180 9. Neurotransmitter Receptors M. NEAL WAXHAM Ionotropic Receptors 181 G-Protein Coupled Receptors 193 References 203 10. Intracellular Signaling HOWARD SCHULMAN AND JAMES L. ROBERTS Signaling Through G-Protein-Linked Receptors 205 Modulation of Neuronal Function by Protein Kinases and Phosphatases 214 Intracellular Signaling Affects Nuclear Gene Expression 222 References 226 Suggested Readings 226 11. Postsynaptic Potentials and Synaptic Integration JOHN H. BYRNE Ionotropic Receptors: Mediators of Fast Excitatory and Inhibitory Synaptic Potentials 227 Metabotropic Receptors: Mediators of Slow Synaptic Potentials 239 Integration of Synaptic Potentials 242 References 245 Suggested Readings 245 12. Complex Information Processing in Dendrites GORDON M. SHEPHERD Strategies for Studying Complex Dendrites 247 Building Principles Step by Step 248 An Axon Places Constraints on Dendritic Processing 249 Dendrodendritic Interactions between Axonal Cells 250 Passive Dendritic Trees Can Perform Complex Computations 251 Separation of Dendritic Fields Enhances Complex Information Processing 252 Distal Dendrites Can be Closely Linked to Axonal Output 253 Depolarizing and Hyperpolarizing Dendritic Conductances Interact Dynamically 255 The Axon Hillock-Initial Segment Encodes Global Output 256 Multiple Impulse Initiation Sites Are under Dynamic Control 256 Retrograde Impulse Spread into Dendrites Can have Many Functions 258
FULL CONTENTS Examples of How Voltage-gated Channels enhanc 16. Neurogenesis and migration Dendritic Information Processing 261 MARIANNE BRONNER-FRASER AND MARY E HATTEN Dendritic Spines Are Multifunctional Microintegrative Units 263 Introduction 351 Summary: The Dendritic Tree as a Complex Information Development of the Peripheral Nervous System 352 Processing System 266 Cell Migration in the CNs 361 References 268 References 372 13. Brain Energy Metabolism endings 37 PIERRE J. MAGISTRETTI 17. Growth Cones and Axon Pathfinding ALEX L KOLODKIN AND MARC TESSIER-LAVIGNE Energy Metabolism of the Brain as a Whole Organ 271 Tight Coupling of Neuronal Activity, Blood Flow, and Growth Cones Are Actively Guided 377 Energy Metabolism 274 Guidance Cues for Developing Axons 380 Energy-Producing and Energy-Consuming Processes in the Guidance Cues and the Control of Cytoskeletal Brain 277 Dynamics 391 Brain Energy Metabolism at the Cellular Level 282 Guidance at the Midline: Changing Responses to Glutamate and Nitrogen Metabolism: a Coordinated Multiple Cues 395 Shuttle Between Astrocytes and Neurons 289 References 399 The Astrocyte-Neuron Metabolic Unit 292 References 292 18. Target Selection, Topographic Maps, and Synapse Formation STEVEN J BURDEN, DENNIS D M OLEARY, AND NERVOUS SYSTEM PETER SCHEIFFELE DEVELOPMENT arget ection Development of the Neuromuscular Synapse 416 14. Neural Induction and Pattern Formation Synapse Formation in the Central Nervous System 426 References 434 ANDREW LUMSDEN AND CHRIS KINTNER Suggested Readings 435 Neural Induction 297 19. Programmed Cell Death and Neurotrophic Early Neural Patterning 303 Factors Regionalization of the Central Nervous System 307 Conclusions 318 RONALD W. OPPENHEIM AND CHRISTOPHER S VON BARTHELD References 319 Cell Death and the neurotrophic Hypothesis 438 15. Cellular determination The Origins of Programmed Cell Death and its WILLIAM A. HARRIS AND VOLKER HARTENSTEIN Occurrence in the devele em438 Origins and Generation of Neuronal Progenitors 321 Functions of Neuronal Programmed Cell Death 444 Spatial and Temporal Coordinates of Neuronal Modes of Cell Death in Developing Neurons 445 Specification 323 The mode of neuronal cell death reflects the The Proneural and Neurogenic genes 326 Activation of Distinct Biochemical and molecular Asymmetric Cell Division and Cell Fate 328 mechanisms 447 Central Neurons and Glia 330 Nerve Growth Factor: The Prototype Target-Derived Sensory Neurons of the Peripheral Nervous System 330 Neuronal Survival Factor 450 The Retina: a Collaboration of Intrinsic and Extrinsic The Neurotrophin Family 452 Cues 336 Neurotrophin Receptors 453 Combinatorial Coding in motor Neurons Secretion and Axonal Transport of Neurotrophins and Determination 342 Pro-Neurotrophins 455 Cells of the Cerebral Cortex 344 Signal Transduction Through TRK Receptors 457 Conclusions 348 Cytokines and Growth Factors have Multiple References 348 Activities 458
FULL CONTENTS ix Examples of How Voltage-gated Channels Enhance Dendritic Information Processing 261 Dendritic Spines Are Multifunctional Microintegrative Units 263 Summary: The Dendritic Tree as a Complex Information Processing System 266 References 268 13. Brain Energy Metabolism PIERRE J. MAGISTRETTI Energy Metabolism of the Brain as a Whole Organ 271 Tight Coupling of Neuronal Activity, Blood Flow, and Energy Metabolism 274 Energy-Producing and Energy-Consuming Processes in the Brain 277 Brain Energy Metabolism at the Cellular Level 282 Glutamate and Nitrogen Metabolism: a Coordinated Shuttle Between Astrocytes and Neurons 289 The Astrocyte-Neuron Metabolic Unit 292 References 292 III NERVOUS SYSTEM DEVELOPMENT 14. Neural Induction and Pattern Formation ANDREW LUMSDEN AND CHRIS KINTNER Neural Induction 297 Early Neural Patterning 303 Regionalization of the Central Nervous System 307 Conclusions 318 References 319 15. Cellular Determination WILLIAM A. HARRIS AND VOLKER HARTENSTEIN Origins and Generation of Neuronal Progenitors 321 Spatial and Temporal Coordinates of Neuronal Specifi cation 323 The Proneural and Neurogenic Genes 326 Asymmetric Cell Division and Cell Fate 328 Central Neurons and Glia 330 Sensory Neurons of the Peripheral Nervous System 330 The Retina: a Collaboration of Intrinsic and Extrinsic Cues 336 Combinatorial Coding in Motor Neurons Determination 342 Cells of the Cerebral Cortex 344 Conclusions 348 References 348 16. Neurogenesis and Migration MARIANNE BRONNER-FRASER AND MARY E. HATTEN Introduction 351 Development of the Peripheral Nervous System 352 Cell Migration in the CNS 361 References 372 Suggested Readings 375 17. Growth Cones and Axon Pathfi nding ALEX L. KOLODKIN AND MARC TESSIER-LAVIGNE Growth Cones Are Actively Guided 377 Guidance Cues for Developing Axons 380 Guidance Cues and the Control of Cytoskeletal Dynamics 391 Guidance at the Midline: Changing Responses to Multiple Cues 395 References 399 18. Target Selection, Topographic Maps, and Synapse Formation STEVEN J. BURDEN, DENNIS D.M. O’LEARY, AND PETER SCHEIFFELE Target Selection 401 Development of the Neuromuscular Synapse 416 Synapse Formation in the Central Nervous System 426 References 434 Suggested Readings 435 19. Programmed Cell Death and Neurotrophic Factors RONALD W. OPPENHEIM AND CHRISTOPHER S. VON BARTHELD Cell Death and the Neurotrophic Hypothesis 438 The Origins of Programmed Cell Death and its Widespread Occurrence in the Developing Nervous System 438 Functions of Neuronal Programmed Cell Death 444 Modes of Cell Death in Developing Neurons 445 The Mode of Neuronal Cell Death Refl ects the Activation of Distinct Biochemical and Molecular Mechanisms 447 Nerve Growth Factor: The Prototype Target-Derived Neuronal Survival Factor 450 The Neurotrophin Family 452 Neurotrophin Receptors 453 Secretion and Axonal Transport of Neurotrophins and Pro-Neurotrophins 455 Signal Transduction Through TRK Receptors 457 Cytokines and Growth Factors have Multiple Activities 458
FULL CONTENTS Programmed Cell Death Is Regulated by Interactions with Peripheral Organization and Processing 540 Targets, Afferents, and Nonneuronal Cells 463 Central Pathways and Processing 542 The Role of Programmed Cell Death in Sensory Cortex 544 464 References 466 References 548 Suggested Readings 467 Suggested Readings 548 20. Synapse Elimination UAN C. TAPIA AND JEFF W. LICHTMAN 24. Chemical Senses: Taste and olfaction Overview 469 KRISTIN SCOTT The Purpose of Synapse Elimination 472 A Structural Analysis of Synapse Elimination at The Taste 549 Neuromuscular Junction 475 Olfaction 560 Role for Interaxonal Competition and Activity 478 Pheromone Detection 576 Is Synapse Elimination Strictly A Developmenta References 578 Summary 488 erences 25. Somatosensor 21. Dendritic Development STEWART HENDRY AND STEVEN HSIAO HOLLIS CLINE, ANIRVAN GHOSH, AND YUH-NUNG JAN Peripheral Mechanisms of Somatic Dynamics of Dendritic Arbor Development 491 Sensation 581 Genetic Control of Dendrite Development Nociception, Thermoreception, and Drosophila 492 Itch 589 Extracellular Regulation of Dendritic Development in Cns Components of Somatic Sensation 592 The Mammalian Brain 496 Thalamic Mechanisms of Somatic Effect of Experience on Dendritic Devel ensation 598 Mechanisms that Mediate Activity-Dependent Dendritic The Path From Nociception to Pain 598 Growth 507 The Trigeminal System 602 Convergence and Divergence 508 Cortical Representation of Touch 604 Conclusion 510 References 607 References 510 Suggested Readings 608 22. Early Experience and Sensitive Periods 26. Audition ERIC I KNUDSEN M. CHRISTIAN BROWN AND JOSEPH SANTOS-SACCHI Birdsong: Learned by Experience 517 External and middle ear 609 Sound Localization: Calibrated by Early Experience in the The Cochlea 610 Owl 520 The Auditory Nerve 618 Principles of Developmental Learning 529 Central Nervous System 624 References 635 Suggested Readings 636 SENSORY SYSTEMS 27. Vision R CLAY REID AND W. MARTIN USREY 23. Fundamentals of Sensory Systems STEWART H. HENDRY. STEVEN S HSIAO, AND Overview 637 M. CHRISTIAN BROWN The Eye and the Retina 639 The Retinogeniculocortical Pathway 649 Sensation and Perception 535 References 658 Receptors 537 Readings 659
x FULL CONTENTS Programmed Cell Death Is Regulated by Interactions with Targets, Afferents, and Nonneuronal Cells 463 The Role of Programmed Cell Death in Neuropathology 464 References 466 Suggested Readings 467 20. Synapse Elimination JUAN C. TAPIA AND JEFF W. LICHTMAN Overview 469 The Purpose of Synapse Elimination 472 A Structural Analysis of Synapse Elimination at The Neuromuscular Junction 475 A Role for Interaxonal Competition and Activity 478 Is Synapse Elimination Strictly A Developmental Phenomenon? 488 Summary 488 References 489 21. Dendritic Development HOLLIS CLINE, ANIRVAN GHOSH, AND YUH-NUNG JAN Dynamics of Dendritic Arbor Development 491 Genetic Control of Dendrite Development in Drosophila 492 Extracellular Regulation of Dendritic Development in The Mammalian Brain 496 Effect of Experience on Dendritic Development 504 Mechanisms that Mediate Activity-Dependent Dendritic Growth 507 Convergence and Divergence 508 Conclusion 510 References 510 22. Early Experience and Sensitive Periods ERIC I. KNUDSEN Birdsong: Learned by Experience 517 Sound Localization: Calibrated by Early Experience in the Owl 520 Principles of Developmental Learning 529 References 532 IV SENSORY SYSTEMS 23. Fundamentals of Sensory Systems STEWART H. HENDRY, STEVEN S. HSIAO, AND M. CHRISTIAN BROWN Sensation and Perception 535 Receptors 537 Peripheral Organization and Processing 540 Central Pathways and Processing 542 Sensory Cortex 544 Summary 548 References 548 Suggested Readings 548 24. Chemical Senses: Taste and Olfaction KRISTIN SCOTT Taste 549 Olfaction 560 Pheromone Detection 576 References 578 Suggested Readings 579 25. Somatosensory System STEWART HENDRY AND STEVEN HSIAO Peripheral Mechanisms of Somatic Sensation 581 Nociception, Thermoreception, and Itch 589 Cns Components of Somatic Sensation 592 Thalamic Mechanisms of Somatic Sensation 598 The Path From Nociception to Pain 598 The Trigeminal System 602 Cortical Representation of Touch 604 References 607 Suggested Readings 608 26. Audition M. CHRISTIAN BROWN AND JOSEPH SANTOS-SACCHI External and Middle Ear 609 The Cochlea 610 The Auditory Nerve 618 Central Nervous System 624 References 635 Suggested Readings 636 27. Vision R. CLAY REID AND W. MARTIN USREY Overview 637 The Eye and the Retina 639 The Retinogeniculocortical Pathway 649 References 658 Suggested Readings 659
FULL CONTENTS 33. Eye movements MOTOR SYSTEMS RICHARD J. KRAUZLIS Eye Movements are Used to Stabilize Gaze or to Shift 28. Fundamentals of Motor Systems Gaze 775 STEN GRILLNER The Mechanics of Moving the Eyes 778 The Fundamental Circuits for Stabilizing Gaze 780 Basic Components of the Motor System 665 The Commands for Shifting gaze are formed in the Motor Programs Coordinate Basic Motor Patterns 667 Brain Stem 782 Roles of Different Parts of the Nervous System in the Gaze Shifts are Controlled by the Midbrain and Control of moy Forebrain 785 Conclusion 676 The Control of Gaze Shifts Involves Higher-Order References 676 Processes 788 The Control of Eye Movements Changes Over Time 790 29. The Spinal and Peripheral Motor System Conclusions 791 MARY KAY FLOETER AND GEORGE Z MENTIS References 792 Suggested Readings 792 Locomotion is a Cycle 677 Connecting the Spinal Cord to the Periphery 680 VI Spinal Interneuron Networks 686 REGULATORY SYSTEMS Descending Control of Spinal Circuits 693 Modulation 693 References 697 34. The Hypothalamus: An Overview of Suggested Readings 697 Regulatory systems J. PATRICK CARD, LARRY W. SWANSON, AND 30. Descending Control of movement ROBERT Y. MOORE MARC H SCHIEBER AND JAMES F. BAKER Historical Perspective 795 The Medial Postural System 699 Hypothalamic Cytoarchitecture 796 The Lateral Voluntary System 710 Functional Organization of the Hypothalamus 797 Effector Systems of the Hypothalamus are Hormonal and References 724 References 805 31. The basal ganglia Suggested readings 806 JONATHAN W MINK 35. Central Control of autonomic functions Basal Ganglia Anatomy 725 Organization of the Autonomic Signaling in basal ganglia 734 Nervous System The Effect of Basal Ganglia Damage on Movement 737 TERRY L POWLEY Fundamental Principles of Basal Ganglia Operation for Motor Control 742 Sympathetic Division: Organized to Mobilize the body for Basal Ganglia Participation in Nonmotor Activity 809 Functions 744 Parasympathetic Division: Organized for Energy References 749 Suggested Readings 750 The Enteric Division of the ans The Nerve net found in the Walls of Visceral Organs 816 32. Cerebellum Ans Pharmacology: Transmitter and Receptor MICHAEL D MAUK AND W. THOMAS THACH Coding 816 Autonomic coordination of homeostasis 819 Anatomy and Phylogenetic Development of the Hierarchically Organized ANS Circuits in the CNS 823 Cerebellum 751 Perspective: Future of the Autonomic Nervous Assessing Cerebellar Function 758 System 825 References 770 ummary and General Conclusions 827
FULL CONTENTS xi V MOTOR SYSTEMS 28. Fundamentals of Motor Systems STEN GRILLNER Basic Components of the Motor System 665 Motor Programs Coordinate Basic Motor Patterns 667 Roles of Different Parts of the Nervous System in the Control of Movement 668 Conclusion 676 References 676 29. The Spinal and Peripheral Motor System MARY KAY FLOETER AND GEORGE Z. MENTIS Locomotion is a Cycle 677 Connecting the Spinal Cord to the Periphery 680 Spinal Interneuron Networks 686 Descending Control of Spinal Circuits 693 Sensory Modulation 693 References 697 Suggested Readings 697 30. Descending Control of Movement MARC H. SCHIEBER AND JAMES F. BAKER The Medial Postural System 699 The Lateral Voluntary System 710 Summary 723 References 724 31. The Basal Ganglia JONATHAN W. MINK Basal Ganglia Anatomy 725 Signaling in Basal Ganglia 734 The Effect of Basal Ganglia Damage on Movement 737 Fundamental Principles of Basal Ganglia Operation for Motor Control 742 Basal Ganglia Participation in Nonmotor Functions 744 References 749 Suggested Readings 750 32. Cerebellum MICHAEL D. MAUK AND W. THOMAS THACH Anatomy and Phylogenetic Development of the Cerebellum 751 Assessing Cerebellar Function 758 References 770 33. Eye Movements RICHARD J. KRAUZLIS Eye Movements are Used to Stabilize Gaze or to Shift Gaze 775 The Mechanics of Moving the Eyes 778 The Fundamental Circuits for Stabilizing Gaze 780 The Commands for Shifting Gaze are Formed in the Brain Stem 782 Gaze Shifts are Controlled by the Midbrain and Forebrain 785 The Control of Gaze Shifts Involves Higher-Order Processes 788 The Control of Eye Movements Changes Over Time 790 Conclusions 791 References 792 Suggested Readings 792 VI REGULATORY SYSTEMS 34. The Hypothalamus: An Overview of Regulatory Systems J. PATRICK CARD, LARRY W. SWANSON, AND ROBERT Y. MOORE Historical Perspective 795 Hypothalamic Cytoarchitecture 796 Functional Organization of the Hypothalamus 797 Effector Systems of the Hypothalamus are Hormonal and Synaptic 800 References 805 Suggested Readings 806 35. Central Control of Autonomic Functions: Organization of the Autonomic Nervous System TERRY L. POWLEY Sympathetic Division: Organized to Mobilize the Body for Activity 809 Parasympathetic Division: Organized for Energy Conservation 812 The Enteric Division of the ANS: The Nerve Net Found in the Walls of Visceral Organs 816 Ans Pharmacology: Transmitter and Receptor Coding 816 Autonomic Coordination of Homeostasis 819 Hierarchically Organized ANS Circuits in the CNS 823 Perspective: Future of the Autonomic Nervous System 825 Summary and General Conclusions 827
FULL CONTENTS References 827 Suggested Readings 828 Suggested Readings 903 36. Neural Regulation of the 40. Neuroendocrine Systems Cardiovascular System ANDREA C. GORE OHN LONGHURST The Hypothalamus Is a Neuroendocrine Organ 905 An Anatomical framework 829 Hypothalamic Releasing/Inhibiting Hormones and their Anatomy and Chemical Properties of Autonomic Targets 906 Pathways 834 The Hypothalamic-Adenohypophysial Neuroendocrine Network Generators 837 Systems 909 Short Term control mechanisms 838 The Hypothalamic-Neurohypophysial Systems 925 Reflex Control of the Cardiovascular System 838 Hormones and the brain 926 Arterial Baroreceptors 838 References 929 Peripheral Arterial Chemoreceptors 844 Cardiac Receptors 847 41. Circadian Timekeeping Abdominal Visceral Reflexes 849 DAVID R WEAVER AND STEVEN M. REPPERT References 852 Overview of the mammalian Circadian timing 37. Neural Control of breathing JACK L FELDMAN AND DONALD R MCCRIMMON The Suprachiasmatic Nuclei Are the Site of the Primary Circadian Pacemaker in Mammals 933 Early Neuroscience and the Brain Stem 85 A Hierarchy of Cell-Autonomous Circadian Central n stem and Breathing 856 Oscillators 934 Where are the Neurons Generating Respiratory The molecular Basis for Circadian Oscillation is a Pattern? 857 Transcriptional Feedback L Discharge Patterns of Respiratory Neurons 859 Circadian Photoreception 943 Where are the Neurons that generate the breathing Rhythm? 862 Diversity of Output Pathways Leading to Physiologi sory Inputs and Altered Breathing 865 Rhythms 950 Mechanoreceptors in the Lungs Adjust Breathing Pattern General Summary 955 and Initiate Protective Reflexes 867 References 956 dulation and Plasticity of Respiratory Motor Suggested Readings 957 Output 868 Suprapontine Structures and Breathing 870 42. Sleep, Dreaming, and Wakefulness EDWARD F. PACE-SCHOTT, J. ALLAN HOBSON AND ROBERT STICKGOLD Food Intake and metabolism The Two States of Sleep: Rapid Eye Movement and STEPHEN C. WOODS AND EDWARD M. STRICKER Nonrapid Eye Movement 959 Sleep in the modern era of Neuroscience 962 Caloric Homeostasis 873 Anatomy and Physiology of Brain Stem Regulatory Role of caloric Homeostasis in Control of Food Systems 965 Intake 875 Modeling the Control of Behavioral State 975 Central Control of food Intake 882 Sleep has Multiple Functions 980 Neuropeptides and the Control of Food Intake 884 References 888 ed Readings 888 Suggested readings 985 43. Reward, Motivation, and Addiction 39. Water Intake and Body fluids GEORGE F KOOB, BARRY J. EVERITT, AND EDWARD M. STRICKER AND JOSEPH G. VERBALIS TREVOR W. ROBBINS Body Fluid Physiology Reward and Motivation 987 smotic ho Addiction 999 References 1014
xii FULL CONTENTS References 827 Suggested Readings 828 36. Neural Regulation of the Cardiovascular System JOHN LONGHURST An Anatomical Framework 829 Anatomy and Chemical Properties of Autonomic Pathways 834 Network Generators 837 Short-Term Control Mechanisms 838 Refl ex Control of the Cardiovascular System 838 Arterial Baroreceptors 838 Peripheral Arterial Chemoreceptors 844 Cardiac Receptors 847 Abdominal Visceral Refl exes 849 References 852 37. Neural Control of Breathing JACK L. FELDMAN AND DONALD R. MCCRIMMON Early Neuroscience and the Brain Stem 855 Central Nervous System and Breathing 856 Where are the Neurons Generating Respiratory Pattern? 857 Discharge Patterns of Respiratory Neurons 859 Where are the Neurons that Generate the Breathing Rhythm? 862 Sensory Inputs and Altered Breathing 865 Mechanoreceptors in the Lungs Adjust Breathing Pattern and Initiate Protective Refl exes 867 Modulation and Plasticity of Respiratory Motor Output 868 Suprapontine Structures and Breathing 870 References 872 38. Food Intake and Metabolism STEPHEN C. WOODS AND EDWARD M. STRICKER Caloric Homeostasis 873 Role of Caloric Homeostasis in Control of Food Intake 875 Central Control of Food Intake 882 Neuropeptides and the Control of Food Intake 884 References 888 Suggested Readings 888 39. Water Intake and Body Fluids EDWARD M. STRICKER AND JOSEPH G. VERBALIS Body Fluid Physiology 889 Osmotic Homeostasis 890 Volume Homeostasis 898 References 902 Suggested Readings 903 40. Neuroendocrine Systems ANDREA C. GORE The Hypothalamus Is a Neuroendocrine Organ 905 Hypothalamic Releasing/Inhibiting Hormones and their Targets 906 The Hypothalamic–Adenohypophysial Neuroendocrine Systems 909 The Hypothalamic-Neurohypophysial Systems 925 Hormones and the Brain 926 References 929 41. Circadian Timekeeping DAVID R. WEAVER AND STEVEN M. REPPERT Overview of the Mammalian Circadian Timing System 931 The Suprachiasmatic Nuclei Are the Site of the Primary Circadian Pacemaker in Mammals 933 A Hierarchy of Cell-Autonomous Circadian Oscillators 934 The Molecular Basis for Circadian Oscillation Is a Transcriptional Feedback Loop 936 Circadian Photoreception 943 Circadian Output Mechanisms 949 Diversity of Output Pathways Leading to Physiological Rhythms 950 General Summary 955 References 956 Suggested Readings 957 42. Sleep, Dreaming, and Wakefulness EDWARD F. PACE-SCHOTT, J. ALLAN HOBSON AND ROBERT STICKGOLD The Two States of Sleep: Rapid Eye Movement and Nonrapid Eye Movement 959 Sleep in the Modern Era of Neuroscience 962 Anatomy and Physiology of Brain Stem Regulatory Systems 965 Modeling the Control of Behavioral State 975 Sleep has Multiple Functions 980 References 982 Suggested Readings 985 43. Reward, Motivation, and Addiction GEORGE F. KOOB, BARRY J. EVERITT, AND TREVOR W. ROBBINS Reward and Motivation 987 Addiction 999 References 1014
FULL CONTENTS VII References 1110 BEHAVIORAL and COGnitive Suggested readings 1111 NEUROSCIENCE 48. Attention JOHN H. REYNOLDS, JACQUELINE P GOTTLIEB, AND SABINE KASTNER 44. Human Brain Evolution JON H. KAAS AND TODD M. PREUSS Introduction 1113 Varieties of atte Evolutionary and Comparative Principles 1019 Neglect Syndrome: A Deficit of Spatial Attention 1114 Evolution of Primate Brains 1027 Single Unit Recording Studies in Nonhuman Primates Why Brain Size Is Important 1034 Provide convergent evidence for a fronto-Parietal Conclusions 1036 Attentional Control System 1116 References 1037 Attention Affects Neural Activity in the Human Visual Suggested Readings 1037 Cortex in the presence and absence of visual Stimulation 1121 45. Cognitive Development and Aging Attention Increases Sensitivity and Boosts the Clarity of PETER R RAPP AND JOCELYNE BACHEVALIER Signals Generated by Neurons in Parts of the Visual System Devoted to Processing Information about velopme Objects 1122 Cognitive Development and Aging: A Life Span Attention Modulates Neural Responses in the Human 1043 ateral geniculate nucleus 1123 Pathological Processes In Cognitive Development and The Visual Search Paradigm has been Used to Study the Aging 1055 Role of Attention in Selecting Relevant Stimuli from eferences Within a Cluttered Visual Environment 1124 Suggested Readings 1066 Where Is the Computational Bottleneck as Revealed by ch Tasks? 1126 46. Visual Perception of Objects Neuronal Receptive Fields Are a Possible Neural Correlate of Limited Capacity 1126 LUIZ PESSOA. ROGER B H. TOOTELL. AND Competition Can Be Biased by Nonspatial LESLIE G. UNGERLEIDER Feedback 1127 Filtering of Unwanted Information in Humans 1129 The Problem of Object Recognition 1067 Conclusions 1130 Substrates for Object Perception and Recognition: Early References 1131 Evidence from Brain damage ted readings Visual Pathways for Object Processing in Nonhum Primates 1071 49. Learning and Memory: Basic Mechanisms Neuronal Properties Within the Object Recognition Pathway 1074 JOHN H. BYRNE Functional Neuroimaging and Electrophysiology of Object Recognition in Humans 1080 Paradigms have been Developed to Study Associative and Nonassociative Learning 1133 Perception and Recognition of Specific Classes of Invertebrate Studies: Key Insights From Aplysia Into Basic Objects 1083 Mechanisms of Learning 1134 Overall Summary 1088 Vertebrate Studies: Long-Term Potentiation 1140 References 1088 1089 Long-Term Depression 1148 Suggested Read How Does a Change in Synaptic Strength Store a Complex Memory? 1149 47. Spatial Cognition References 1151 CAROL L. COLBY AND CARL R. OLSON Suggested Readings 1152 Neural Systems for Spatial Cognition 1091 50. Learning and Memory: Brain Systems Parietal Cortex 1092 JOSEPH R MANNS AND HOW ARD EICHENBAUM Frontal cortex 1102 Hippocampus and Adjacent Cortex 1107 Introduction 1153 Spatial Cognition and Spatial Action 1109 Hi of me
FULL CONTENTS xiii VII BEHAVIORAL and COGNITIVE NEUROSCIENCE 44. Human Brain Evolution JON H. KAAS AND TODD M. PREUSS Evolutionary and Comparative Principles 1019 Evolution of Primate Brains 1027 Why Brain Size Is Important 1034 Conclusions 1036 References 1037 Suggested Readings 1037 45. Cognitive Development and Aging PETER R. RAPP AND JOCELYNE BACHEVALIER Brain Development 1039 Cognitive Development and Aging: A Life Span Perspective 1043 Pathological Processes In Cognitive Development and Aging 1055 References 1065 Suggested Readings 1066 46. Visual Perception of Objects LUIZ PESSOA, ROGER B. H. TOOTELL, AND LESLIE G. UNGERLEIDER The Problem of Object Recognition 1067 Substrates for Object Perception and Recognition: Early Evidence from Brain Damage 1068 Visual Pathways for Object Processing in Nonhuman Primates 1071 Neuronal Properties Within the Object Recognition Pathway 1074 Functional Neuroimaging and Electrophysiology of Object Recognition in Humans 1080 Perception and Recognition of Specifi c Classes of Objects 1083 Overall Summary 1088 References 1088 Suggested Readings 1089 47. Spatial Cognition CAROL L. COLBY AND CARL R. OLSON Neural Systems for Spatial Cognition 1091 Parietal Cortex 1092 Frontal Cortex 1102 Hippocampus and Adjacent Cortex 1107 Spatial Cognition and Spatial Action 1109 References 1110 Suggested Readings 1111 48. Attention JOHN H. REYNOLDS, JACQUELINE P. GOTTLIEB, AND SABINE KASTNER Introduction 1113 Varieties of Attention 1113 Neglect Syndrome: A Defi cit of Spatial Attention 1114 Single Unit Recording Studies in Nonhuman Primates Provide Convergent Evidence for A Fronto-Parietal Attentional Control System 1116 Attention Affects Neural Activity in the Human Visual Cortex in the Presence and Absence of Visual Stimulation 1121 Attention Increases Sensitivity and Boosts the Clarity of Signals Generated by Neurons in Parts of the Visual System Devoted to Processing Information about Objects 1122 Attention Modulates Neural Responses in the Human Lateral Geniculate Nucleus 1123 The Visual Search Paradigm has been Used to Study the Role of Attention in Selecting Relevant Stimuli from Within a Cluttered Visual Environment 1124 Where Is the Computational Bottleneck as Revealed by Search Tasks? 1126 Neuronal Receptive Fields Are a Possible Neural Correlate of Limited Capacity 1126 Competition Can Be Biased by Nonspatial Feedback 1127 Filtering of Unwanted Information in Humans 1129 Conclusions 1130 References 1131 Suggested Readings 1131 49. Learning and Memory: Basic Mechanisms JOHN H. BYRNE Paradigms have been Developed to Study Associative and Nonassociative Learning 1133 Invertebrate Studies: Key Insights From Aplysia Into Basic Mechanisms of Learning 1134 Vertebrate Studies: Long-Term Potentiation 1140 Long-Term Depression 1148 How Does a Change in Synaptic Strength Store a Complex Memory? 1149 References 1151 Suggested Readings 1152 50. Learning and Memory: Brain Systems JOSEPH R. MANNS AND HOWARD EICHENBAUM Introduction 1153 History of Memory Systems 1153