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SINAUER ASSOCIATES, INC. • Publishers Sunderland, Massachusetts U.S.A. © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTENTS IN BRIEF 1 Studying the Nervous System 1 Unit I Neural Signaling 2 3 4 5 6 7 8 Electrical Signals of Nerve Cells 25 Voltage-Dependent Membrane Permeability 41 Ion Channels and Transporters 57 Synaptic Transmission 77 Neurotransmitters and Their Receptors 109 Molecular Signaling within Neurons 141 Synaptic Plasticity 163 Unit II Sensation and Sensory Processing 9 10 11 12 13 14 15 The Somatic Sensory System: Touch and Proprioception 189 Pain 209 Vision: The Eye 229 Central Visual Pathways 257 The Auditory System 277 The Vestibular System 303 The Chemical Senses 321 Unit III Movement and Its Central Control 16 17 18 19 20 21 Lower Motor Neuron Circuits and Motor Control 353 Upper Motor Neuron Control of the Brainstem and Spinal Cord 375 Modulation of Movement by the Basal Ganglia 399 Modulation of Movement by the Cerebellum 417 Eye Movements and Sensory Motor Integration 435 The Visceral Motor System 451 Unit IV The Changing Brain 22 23 24 25 Early Brain Development 477 Construction of Neural Circuits 507 Modification of Neural Circuits as a Result of Experience 537 Repair and Regeneration in the Nervous System 559 Unit V Complex Brain Functions 26 27 28 29 30 31 Association Cortex and Cognition 587 Speech and Language 607 Sleep and Wakefulness 625 Emotions 647 Sex, Sexuality, and the Brain 669 Memory 695 Appendix: Survey of Human Neuroanatomy 717 Atlas: The Human Central Nervous System 745 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTENTS CHAPTER 1 Studying the Nervous System 1 Overview 1 Genetics, Genomics, and the Brain 1 BOX 1A MODEL ORGANISMS IN NEUROSCIENCE 2 The Cellular Components of the Nervous System 4 Neurons 6 Glial Cells 7 Cellular Diversity in the Nervous System 10 Neural Circuits 10 The Organization of the Human Nervous System 13 Neural Systems 15 Structural Analysis of Neural Systems 15 Functional Analysis of Neural Systems 16 BOX 1B BRAIN IMAGING TECHNIQUES 18 Analyzing Complex Behavior 21 Summary 21 Additional Reading 21 UNIT I NEURAL SIGNALING 23 Summary 40 Additional Reading 40 CHAPTER 3 CHAPTER 2 Electrical Signals of Nerve Cells 25 Overview 25 Electrical Signals of Nerve Cells 25 Long-Distance Transmission of Electrical Signals 27 How Ion Movements Produce Electrical Signals 29 BOX 2A PASSIVE MEMBRANE PROPERTIES 30 Forces That Create Membrane Potentials 33 Electrochemical Equilibrium in an Environment with More Than One Permeant Ion 34 The Ionic Basis of the Resting Membrane Potential 35 BOX 2B THE REMARKABLE GIANT NERVE CELLS OF SQUID 36 The Ionic Basis of Action Potentials 37 BOX 2C ACTION POTENTIAL FORM AND NOMENCLATURE 39 Voltage-Dependent Membrane Permeability 41 Overview 41 Ionic Currents across Nerve Cell Membranes 41 BOX 3A THE VOLTAGE CLAMP METHOD 42 Two Types of Voltage-Dependent Ionic Currents 43 Two Voltage-Dependent Membrane Conductances 45 Reconstruction of the Action Potential 47 BOX 3B THRESHOLD 49 Long-Distance Signaling by Means of Action Potentials 51 Increased Conduction Velocity as a Result of Myelination 51 Summary 53 BOX 3C MULTIPLE SCLEROSIS 54 Additional Reading 55 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. VIII CONTENTS CHAPTER 4 CHAPTER 6 Ion Channels and Transporters 57 Neurotransmitters and Their Receptors 109 Overview 57 Ion Channels Underlying Action Potentials 57 BOX 4A THE PATCH CLAMP METHOD 59 BOX 4B TOXINS THAT POISON ION CHANNELS 62 The Diversity of Ion Channels 63 BOX 4C EXPRESSION OF ION CHANNELS IN XENOPUS OOCYTES 63 Voltage-Gated Ion Channels 64 Ligand-Gated Ion Channels 66 Stretch- and Heat-Activated Channels 66 The Molecular Structure of Ion Channels 66 Active Transporters Create and Maintain Ion Gradients 69 BOX 4D DISEASES CAUSED BY ALTERED ION CHANNELS 70 Functional Properties of the Na+/K+ Pump 72 The Molecular Structure of ATPase Pumps 72 Summary 74 Additional Reading 75 CHAPTER 5 Synaptic Transmission 77 Overview 77 Electrical Synapses 78 Signal Transmission at Chemical Synapses 80 Properties of Neurotransmitters 81 BOX 5A CRITERIA THAT DEFINE A NEUROTRANSMITTER 84 Quantal Release of Neurotransmitters 84 Release of Transmitters from Synaptic Vesicles 86 Local Recycling of Synaptic Vesicles 86 The Role of Calcium in Transmitter Secretion 88 Molecular Mechanisms of Synaptic Vesicle Cycling 90 BOX 5B DISEASES THAT AFFECT THE PRESYNAPTIC TERMINAL 93 Neurotransmitter Receptors 96 Postsynaptic Membrane Permeability Changes during Synaptic Transmission 97 Excitatory and Inhibitory Postsynaptic Potentials 101 Summation of Synaptic Potentials 102 BOX 5C THE “TRIPARTITE SYNAPSE” 104 Summary 105 Additional Reading 106 Overview 109 Categories of Neurotransmitters 109 Acetylcholine 111 BOX 6A NEUROTOXINS THAT ACT ON POSTSYNAPTIC RECEPTORS 114 Glutamate 116 BOX 6B MYASTHENIA GRAVIS: AN AUTOIMMUNE DISEASE OF NEUROMUSCULAR SYNAPSES 117 BOX 6C EXCITOTOXICITY FOLLOWING ACUTE BRAIN INJURY 121 GABA and Glycine 122 BOX 6D EXCITATORY ACTIONS OF GABA IN THE DEVELOPING BRAIN 124 The Biogenic Amines 125 BOX 6E BIOGENIC AMINE NEUROTRANSMITTERS AND PSYCHIATRIC DISORDERS 126 BOX 6F ADDICTION 128 ATP and Other Purines 131 Peptide Neurotransmitters 132 Unconventional Neurotransmitters 135 BOX 6G MARIJUANA AND THE BRAIN 137 Summary 139 Additional Reading 139 CHAPTER 7 Molecular Signaling within Neurons 141 Overview 141 Strategies of Molecular Signaling 141 The Activation of Signaling Pathways 143 Receptor Types 144 G-Proteins and Their Molecular Targets 145 Second Messengers 147 BOX 7A DYNAMIC IMAGING OF INTRACELLULAR SIGNALING 149 Second Messenger Targets: Protein Kinases and Phosphatases 151 Nuclear Signaling 153 BOX 7B DENDRITIC SPINES 154 Examples of Neuronal Signal Transduction 157 Summary 159 Additional Reading 160 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTE NTS CHAPTER 8 IX Mechanisms Underlying LTP 173 Synaptic Plasticity 163 Overview 163 Short-Term Synaptic Plasticity 163 Long-Term Synaptic Plasticity Underlies Behavioral Modification in Aplysia 166 Long-Term Potentiation at a Hippocampal Synapse 169 BOX 8A GENETICS OF LEARNING AND MEMORY IN THE FRUIT FLY 170 BOX 8B SILENT SYNAPSES 175 Mechanisms Underlying LTD 177 Spike-Timing Dependent Plasticity 181 BOX 8C EPILEPSY: THE EFFECT OF PATHOLOGICAL ACTIVITY ON NEURAL CIRCUITRY 182 Summary 183 Additional Reading 184 UNIT II SENSATION AND SENSORY PROCESSING 187 Beyond SI: Corticocortical and Descending Pathways 205 Plasticity in the Adult Cerebral Cortex 206 Summary 207 Additional Reading 207 CHAPTER 9 The Somatic Sensory System: Touch and Proprioception 189 Overview 189 Afferent Fibers Convey Somatic Sensory Information to the Central Nervous System 189 Somatic Sensory Afferents Exhibit Distinct Functional Properties 191 BOX 9A DERMATOMES 191 Mechanoreceptors Specialized to Receive Tactile Information 194 Mechanoreceptors Specialized for Proprioception 196 Central Pathways Conveying Tactile Information from the Body: The Dorsal Column–Medial Lemniscal System 198 Central Pathways Conveying Tactile Information from the Face: The Trigeminothalamic System 200 Central Pathways Conveying Proprioceptive Information from the Body 200 Central Pathways Conveying Proprioceptive Information from the Face 201 The Somatic Sensory Components of the Thalamus 201 Primary Somatic Sensory Cortex 202 BOX 9B PATTERNS OF ORGANIZATION WITHIN THE SENSORY CORTICES: BRAIN MODULES 203 CHAPTER 10 Pain 209 Overview 209 Nociceptors 209 Transduction and Transmission of Nociceptive Signals 211 BOX 10A CAPSAICIN 212 Central Pain Pathways Are Distinct from Mechanosensory Pathways 213 BOX 10B REFERRED PAIN 214 BOX 10C A DORSAL COLUMN PATHWAY FOR VISCERAL PAIN 215 Parallel Pain Pathways 217 Pain and Temperature Pathways for the Face 218 Other Modalities Mediated by the Anterolateral System 220 Sensitization 220 Descending Control of Pain Perception 223 BOX 10D PHANTOM LIMBS AND PHANTOM PAIN 222 The Placebo Effect 223 The Physiological Basis of Pain Modulation 224 Summary 226 Additional Reading 226 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. X CONTENTS CHAPTER 13 CHAPTER 11 Vision: The Eye 229 Overview 229 Anatomy of the Eye 229 The Formation of Images on the Retina 230 BOX 11A MYOPIA AND OTHER REFRACTIVE ERRORS 231 The Surface of the Retina 233 Retinal Circuitry 233 BOX 11B THE BLIND SPOT 234 BOX 11C MACULAR DEGENERATION 235 BOX 11D RETINITIS PIGMENTOSA 237 Retinal Pigment Epithelium 238 Phototransduction 238 Functional Specialization of the Rod and Cone Systems 242 Anatomical Distribution of Rods and Cones 244 Cones and Color Vision 245 BOX 11E THE IMPORTANCE OF CONTEXT IN COLOR PERCEPTION 246 Retinal Circuits for Detecting Luminance Change 249 BOX 11F THE PERCEPTION OF LIGHT INTENSITY 250 Contribution of Retinal Circuits to Light Adaptation 253 Summary 255 Additional Reading 256 CHAPTER 12 Central Visual Pathways 257 Overview 257 Central Projections of Retinal Ganglion Cells 257 The Retinotopic Representation of the Visual Field 259 Visual Field Deficits 261 Spatiotemporal Tuning Properties of Neurons in Primary Visual Cortex 263 Primary Visual Cortex Architecture 265 Combining Inputs from Two Eyes 267 Division of Labor within the Primary Visual Pathway 269 BOX 12A RANDOM DOT STEREOGRAMS AND RELATED AMUSEMENTS 270 The Functional Organization of Extrastriate Visual Areas 272 Summary 274 Additional Reading 275 The Auditory System 277 Overview 277 Sound 277 The Audible Spectrum 278 A Synopsis of Auditory Function 279 BOX 13A FOUR CAUSES OF ACQUIRED HEARING LOSS 279 BOX 13B MUSIC 280 The External Ear 281 The Middle Ear 282 BOX 13C SENSORINEURAL HEARING LOSS AND COCHLEAR IMPLANTS 283 The Inner Ear 284 Hair Cells and the Mechanoelectrical Transduction of Sound Waves 287 The Ionic Basis of Mechanotransduction in Hair Cells 290 The Cochlear Amplifier 291 Tuning and Timing in the Auditory Nerve 291 BOX 13D THE SWEET SOUND OF DISTORTION 292 How Information from the Cochlea Reaches Targets in the Brainstem 293 Integrating Information from the Two Ears 294 Monaural Pathways from the Cochlear Nucleus to the Nuclei of the Lateral Lemniscus 297 Integration in the Inferior Colliculus 297 The Auditory Thalamus 297 The Auditory Cortex 298 BOX 13E REPRESENTING COMPLEX SOUNDS IN THE BRAINS OF BATS AND HUMANS 299 Summary 301 Additional Reading 301 CHAPTER 14 The Vestibular System 303 Overview 303 The Vestibular Labyrinth 303 BOX 14A A PRIMER ON TERMINOLOGY 304 Vestibular Hair Cells 305 BOX 14B ADAPTATION AND TUNING OF VESTIBULAR HAIR CELLS 306 The Otolith Organs: The Utricle and Saccule 307 How Otolith Neurons Sense Tilts and Linear Accelerations of the Head 309 The Semicircular Canals 310 How Semicircular Canal Neurons Sense Angular Accelerations 312 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTE NTS Central Pathways for Stabilizing Gaze, Head, and Posture 312 BOX 14C CLINICAL EVALUATION OF THE VESTIBULAR SYSTEM 314 BOX 14D MAUTHNER CELLS IN FISH 316 Vestibular Pathways to the Thalamus and Cortex 318 Spatial Orientation Perception and Multisensory Integration 318 Summary 319 Additional Reading 320 CHAPTER 15 The Chemical Senses 321 Overview 321 The Organization of the Olfactory System 321 Olfactory Perception in Humans 323 Assessing Olfactory Function in the Laboratory or Clinic 324 XI BOX 15A YOU ONLY GET ONE NOSE 326 Physiological and Behavioral Responses to Odorants 327 BOX 15B PHEROMONES, REPRODUCTION, AND THE VOMERONASAL SYSTEM 328 Olfactory Epithelium and Olfactory Receptor Neurons 329 Odor Transduction and Odorant Receptor Proteins 332 BOX 15C THE “DOGTOR” IS IN 333 Physiological Mechanisms of Odor Transduction 334 The Olfactory Bulb 336 Cortical Processing of Information Relayed from the Olfactory Bulb 340 The Organization of the Taste System 341 Taste Perception in Humans 343 Taste Receptor Proteins and Transduction 345 Neural Coding in the Taste System 347 Trigeminal Chemoreception 349 Summary 349 Additional Reading 350 UNIT III MOVEMENT AND ITS CENTRAL CONTROL 351 BOX 16B LOCOMOTION IN THE LEECH AND THE LAMPREY 368 Spinal Cord Circuitry and Locomotion 369 BOX 16C THE AUTONOMY OF CENTRAL PATTERN GENERATORS: EVIDENCE FROM THE LOBSTER STOMATOGASTRIC GANGLION 370 CHAPTER 16 Lower Motor Neuron Circuits and Motor Control 353 Overview 353 Neural Centers Responsible for Movement 353 Motor Neuron–Muscle Relationships 355 The Motor Unit 357 The Regulation of Muscle Force 358 BOX 16A MOTOR UNIT PLASTICITY 359 The Spinal Cord Circuitry Underlying Muscle Stretch Reflexes 362 The Influence of Sensory Activity on Motor Behavior 364 Other Sensory Feedback Affecting Motor Performance 365 Flexion Reflex Pathways 367 The Lower Motor Neuron Syndrome 372 BOX 16D AMYOTROPHIC LATERAL SCLEROSIS 373 Summary 373 Additional Reading 374 CHAPTER 17 Upper Motor Neuron Control of the Brainstem and Spinal Cord 375 Overview 375 Organization of Descending Motor Control 375 The Corticospinal and Corticobulbar Tracts 377 BOX 17A PATTERNS OF FACIAL WEAKNESS AND THEIR IMPORTANCE FOR LOCALIZING NEUROLOGICAL INJURY 379 Functional Organization of the Primary Motor Cortex 380 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. XII CONTENTS BOX 17B WHAT DO MOTOR MAPS REPRESENT? 381 BOX 17C SENSORY MOTOR TALENTS AND CORTICAL SPACE 384 The Premotor Cortex 387 Motor Control Centers in the Brainstem: Upper Motor Neurons that Maintain Balance, Govern Posture, and Orient Gaze 389 BOX 17D THE RETICULAR FORMATION 391 Damage to Descending Motor Pathways: The Upper Motor Neuron Syndrome 395 BOX 17E MUSCLE TONE 397 Summary 397 Additional Reading 398 BOX 19B GENETIC ANALYSIS OF CEREBELLAR FUNCTION 431 Summary 433 Additional Reading 433 CHAPTER 20 Eye Movements and Sensory Motor Integration 435 Overview 435 What Eye Movements Accomplish 435 The Actions and Innervation of Extraocular Muscles 436 BOX 20A THE PERCEPTION OF STABILIZED RETINAL IMAGES 437 CHAPTER 18 Modulation of Movement by the Basal Ganglia 399 Overview 399 Projections to the Basal Ganglia 399 Projections from the Basal Ganglia to Other Brain Regions 402 Evidence from Studies of Eye Movements 404 Circuits within the Basal Ganglia System 405 Dopamine Modulates Basal Ganglia Circuits 407 Hypokinetic and Hyperkinetic Movement Disorders 408 BOX 18A PARKINSON’S DISEASE: AN OPPORTUNITY FOR NOVEL THERAPEUTIC APPROACHES 410 BOX 18B HUNTINGTON’S DISEASE 411 BOX 18C DEEP BRAIN STIMULATION 412 BOX 18D BASAL GANGLIA LOOPS AND NON-MOTOR BRAIN FUNCTIONS 414 Types of Eye Movements and Their Functions 438 Neural Control of Saccadic Eye Movements 440 BOX 20B SENSORY MOTOR INTEGRATION IN THE SUPERIOR COLLICULUS 443 BOX 20C FROM PLACE CODES TO RATE CODES 445 Neural Control of Smooth Pursuit Movements 449 Neural Control of Vergence Movements 449 Summary 450 Additional Reading 450 CHAPTER 21 The Visceral Motor System 451 Overview 451 Early Studies of the Visceral Motor System 454 Distinctive Features of the Visceral Motor System 454 The Sympathetic Division of the Visceral Motor System 456 BOX 21A THE HYPOTHALAMUS 456 Summary 416 Additional Reading 416 CHAPTER 19 Modulation of Movement by the Cerebellum 417 Overview 417 Organization of the Cerebellum 417 Projections to the Cerebellum 419 Projections from the Cerebellum 421 Circuits within the Cerebellum 423 Cerebellar Circuitry and the Coordination of Ongoing Movement 426 BOX 19A PRION DISEASES 427 The Parasympathetic Division of the Visceral Motor System 460 The Enteric Nervous System 461 Sensory Components of the Visceral Motor System 462 Central Control of Visceral Motor Functions 464 BOX 21B HORNER’S SYNDROME 465 Neurotransmission in the Visceral Motor System 466 BOX 21C OBESITY AND THE BRAIN 466 Autonomic Regulation of Cardiovascular Function 468 Autonomic Regulation of the Bladder 470 Autonomic Regulation of Sexual Function 472 Summary 473 Additional Reading 474 Further Consequences of Cerebellar Lesions 429 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTE NTS XIII UNIT IV THE CHANGING BRAIN 475 The Axon Growth Cone 509 The Molecular Basis of Growth Cone Motility 510 BOX 23A CHOOSING SIDES: AXON GUIDANCE AT THE OPTIC CHIASM 512 Non-Diffusible Signals for Axon Guidance 513 CHAPTER 22 BOX 23B HUMAN AXON GUIDANCE DISORDERS 515 Early Brain Development 477 Overview 477 Formation of the Nervous System: Gastrulation and Neurulation 477 BOX 22A STEM CELLS: PROMISE AND PERIL 479 Formation of the Major Brain Subdivisions 481 BOX 22B RHOMBOMERES 484 The Molecular Basis of Neural Induction 485 BOX 22C RETINOIC ACID: TERATOGEN AND INDUCTIVE SIGNAL 488 Integrated Inductive Signals Establish Neuron Identity 489 The Initial Differentiation of Neurons and Glia 489 BOX 22D NEUROGENESIS: WHERE, WHEN, AND WHAT 492 Molecular Regulation of Neurogenesis 493 The Generation of Neuronal Diversity 495 Molecular and Genetic Disruptions of Early Neural Development 496 BOX 22E TRIPLE JEOPARDY: DISEASES ASSOCIATED WITH SONIC HEDGEHOG 497 Neuronal Migration in the Peripheral Nervous System 499 BOX 22F MIXING IT UP: LONG-DISTANCE NEURONAL MIGRATION 501 Neuronal Migration in the Central Nervous System 502 Molecular Mechanisms of Neuronal Migration and Cortical Migration Disorders 503 Summary 503 Additional Reading 505 Chemoattraction and Chemorepulsion 517 The Formation of Topographic Maps 519 Selective Synapse Formation 521 Regulation of Neuronal Connections by Trophic Interactions 523 Competitive Interactions and the Formation of Neuronal Connections 525 BOX 23C WHY DO NEURONS HAVE DENDRITES? 528 The Molecular Basis of Trophic Interactions 529 BOX 23D THE DISCOVERY OF BDNF AND THE NEUROTROPHIN FAMILY 530 Neurotrophin Signaling 533 Summary 535 Additional Reading 535 CHAPTER 24 Modification of Neural Circuits as a Result of Experience 537 Overview 537 Neural Activity and Brain Development 537 Critical Periods 538 BOX 24A BUILT-IN BEHAVIORS 540 BOX 24B BIRDSONG 541 Cellular and Molecular Correlates of Activity-Dependent Plasticity during Critical Periods 542 Critical Periods in Visual System Development 543 BOX 24C TRANSNEURONAL LABELING WITH RADIOACTIVE AMINO ACIDS 544 Effects of Visual Deprivation on Ocular Dominance 545 BOX 24D CORRELATION AS CAUSATION: LESSONS FROM A THREE-EYED FROG 549 CHAPTER 23 Construction of Neural Circuits 507 Overview 507 Neuronal Polarization: The First Step in Neural Circuit Formation 507 Manipulating Competition 550 Amblyopia, Strabismus, and Critical Periods for Human Vision 551 Evidence for Critical Periods in Other Sensory Systems 552 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. XIV CONTENTS The Development of Language: A Critical Period for a Distinctly Human Behavior 553 Human Brain Development, Activity-Dependent Plasticity, and Critical Periods 554 Summary 556 Additional Reading 557 Regeneration of Peripheral Synapses 567 BOX 25A SPECIFIC REGENERATION OF SYNAPTIC CONNECTIONS IN AUTONOMIC GANGLIA 568 Regeneration after Damage to the Central Nervous System 570 Cellular and Molecular Responses to Injury 570 BOX 25B CASUALTIES OF WAR AND SPORTS 571 CHAPTER 25 Repair and Regeneration in the Nervous System 559 Overview 559 The Damaged Brain 559 Functional Reorganization without Repair 560 Three Types of Neuronal Repair 561 Peripheral Nerve Regeneration 563 The Cellular and Molecular Basis of Peripheral Nerve Repair 564 Axon Growth after Brain Injury 574 Neurogenesis in the Mature Central Nervous System 575 Adult Neurogenesis in Non-Mammalian Vertebrates 576 Neurogenesis in the Adult Mammalian Brain 577 Cellular and Molecular Mechanisms of Adult Neurogenesis 578 Adult Neurogenesis, Stem Cells, and Brain Repair in Humans 580 BOX 25C NUCLEAR WEAPONS AND NEUROGENESIS 581 Summary 583 Additional Reading 583 UNIT V COMPLEX BRAIN FUNCTIONS 585 “Planning Neurons” in the Monkey Frontal Cortex 603 Summary 605 Additional Reading 605 CHAPTER 27 CHAPTER 26 Association Cortex and Cognition 587 Overview 587 The Association Cortices 587 An Overview of Cortical Structure 588 Unique Features of the Association Cortices 589 BOX 26A CORTICAL LAMINATION 590 The Parietal Association Cortex Mediates Attention 591 “Attention Neurons” in the Monkey Parietal Cortex 594 The Temporal Association Cortex Mediates Recognition 596 “Recognition Neurons” in the Monkey Temporal Cortex 598 The Frontal Association Cortex Mediates Planning and Decision Making 599 BOX 26B NEUROPSYCHOLOGICAL TESTING 600 BOX 26C PSYCHOSURGERY 602 Speech and Language 607 Overview 607 Language Is Localized and Lateralized in the Brain 607 BOX 27A THE GENERATION OF SPEECH 608 Aphasias 610 BOX 27B DO OTHER ANIMALS HAVE LANGUAGE? 611 BOX 27C WORDS AND MEANING 613 Confirmation of Language Lateralization and Other Insights 615 BOX 27D LANGUAGE AND HANDEDNESS 617 The Search for Anatomical Differences between the Right and Left Hemispheres 618 Mapping Language Functions 619 The Role of the Right Hemisphere 622 Sign Language 622 Summary 623 Additional Reading 623 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. CONTE NTS XV CHAPTER 30 CHAPTER 28 Sleep and Wakefulness 625 Overview 625 Why Do Humans (and Many Other Animals) Sleep? 625 BOX 28A THE SLEEP STYLES OF DIFFERENT SPECIES 627 The Circadian Cycle of Sleep and Wakefulness 628 BOX 28B MOLECULAR MECHANISMS OF BIOLOGICAL CLOCKS 629 Stages of Sleep 631 BOX 28C ELECTROENCEPHALOGRAPHY 632 Physiological Changes in Sleep States 635 Other Possible Functions of Sleep and Dreaming 636 Neural Circuits Governing Sleep 637 BOX 28D CONSCIOUSNESS 638 Sex, Sexuality, and the Brain 669 Overview 669 Sexual Dimorphisms and Sexually Dimorphic Behaviors 669 Sex, Gonads, Bodies, and Brains 671 BOX 30A THE SCIENCE OF LOVE (OR, LOVE IS A DRUG) 672 Hormonal Influences on Sexual Dimorphism 674 Primary Sexual Dimorphisms in the Brain 676 Brain Dimorphisms and the Control of Reproductive Behaviors 678 Structural and Functional Dimorphisms for Pregnancy and Parenting 680 BOX 30B THE GOOD MOTHER 683 Emotions 647 Cellular and Molecular Basis of Sexually Dimorphic Structures and Behaviors 684 Steroid Receptors and Responses in the Adult Brain 686 Human Genetic Disorders of Genotypic and Phenotypic Sex 687 Sexual Orientation and the Brain: Molecular and Genetic Analysis 688 Sexual Orientation and Human Brain Structure 690 Sex-Based Differences in Cognitive Functions 691 Summary 693 Additional Reading 693 Overview 647 Physiological Changes Associated with Emotion 647 CHAPTER 31 Thalamocortical Interactions in Sleep 641 BOX 28E DRUGS AND SLEEP 641 Sleep Disorders 643 BOX 28F CHRONIC FATIGUE SYNDROME 644 Summary 645 Additional Reading 645 CHAPTER 29 BOX 29A FACIAL EXPRESSIONS: PYRAMIDAL AND EXTRAPYRAMIDAL CONTRIBUTIONS 648 The Integration of Emotional Behavior 650 The Limbic System 652 The Importance of the Amygdala 653 BOX 29B THE ANATOMY OF THE AMYGDALA 654 BOX 29C THE REASONING BEHIND AN IMPORTANT DISCOVERY 655 BOX 29D FEAR AND THE HUMAN AMYGDALA: A CASE STUDY 657 BOX 29E AFFECTIVE DISORDERS 659 The Relationship between Neocortex and Amygdala 660 Cortical Lateralization of Emotional Functions 661 Emotion, Reason, and Social Behavior 662 Emotional Reinforcement and Addiction 663 Summary 666 Additional Reading 667 Memory 695 Overview 695 Qualitative Categories of Human Memory 695 Temporal Categories of Memory 696 BOX 31A PHYLOGENETIC MEMORY 697 Memory Consolidation and Priming 698 The Importance of Association in Information Storage 698 Conditioned Learning 699 BOX 31B SAVANT SYNDROME 700 Forgetting 702 Brain Systems Underlying Declarative Memory Acquisition and Storage 703 BOX 31C CLINICAL CASES REVEAL AN ANATOMICAL SUBSTRATE FOR DECLARATIVE MEMORIES 703 Brain Systems Underlying Nondeclarative Memory Acquisition and Storage 711 Memory and Aging 712 BOX 31D ALZHEIMER’S DISEASE 713 Summary 714 Additional Reading 715 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. XVI CONTENTS APPENDIX ATLAS Survey of Human Neuroanatomy 717 The Human Central Nervous System 745 Overview 717 Neuroanatomical Terminology 717 Basic Subdivisions of the Central Nervous System 718 External Anatomy of the Spinal Cord 720 Internal Anatomy of the Spinal Cord 721 Brainstem and Cranial Nerves 722 Lateral Surface of the Brain 728 Dorsal and Ventral Surfaces of the Brain 729 Midsagittal Surface of the Brain 730 Internal Anatomy of the Forebrain 731 Plate 1: Brain Surface 746 Plate 2: Coronal MR 748 Plate 3: Axial MR 750 Plate 4: Sagittal MR 754 Plate 5: Brainstem 756 Plate 6: Spinal Cord 758 GLOSSARY G–1 ILLUSTRATION CREDITS IC–1 INDEX I–1 BOX A THALAMUS AND THALAMOCORTICAL RELATIONS 732 Blood Supply of the Brain and Spinal Cord 735 BOX B STROKE 735 The Blood–Brain Barrier 741 The Meninges 742 The Ventricular System 742 References 744 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.