Neuroscience - 7th Edition

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succeed in your neuroscience course
. .. Sy tv i US c. bys. Mark Williams, Leonard E. White, and Andrew Mace

An Interactive Atlas and Visual Glossary of Human Neuroanatomy

Each new copy of Neuroscience, Fifth Edition includes an access code for Sylvius-a unique digital
learning environment for exploring and understanding the structure of the human central nervous
system. Sylvius features fully annotated surface views of the human brain, as well as interactive
tools for dissecting the central nervous system and viewing fully annotated cross-sections of pre-
served specimens and living subjects imaged by magnetic resonance. This newly expanded and re-
configured Sylvfus is more than a conventional atlas; it incorporates a comprehensive, visually-rich,
searchable database of more than 500 neuroanatomical terms that are concisely defined and visual-
ized in photographs, magnetic resonance images, and illustrations from Neuroscience, Fifth Edition.
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• Surface Anatomy Atlases: Photographic, Magnetic Resonance Image, Brainstem Model
• Sectional Anatomy Atlases: Photographic, Magnetic Resonance Image, Brainstem and
Spinal Cord
• Pathways: Allow you to follow the flow of information in several important long-tract pathways
of the central nervous system
(Features and access Instructions continued on next page)
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• Vasual Glossary: Searchable glossary that provides visual :representations, concise anatomical
and functional definitions, and audio pronunciation of neuroanatomical structw:es
• Over 500 neuroanatomical structures and terms are identified and described
• categories of anatomical structures and terms (e.g., cranial nerves, spinal cord tracts, lobes,
cortical areas, etc.) that can be browsed easily
• A notes feature that allows you to type and save notes for any selected structure
• A self-quiz mode that allows assessment of structure identification and functional information

Access Instructions
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SINAUER ASSOCIATES, INC. • Publishers
Sunderland, Massachusetts U.S.A.
THE COVERS
The image on the front cover shows a portion of the two hemispheres of the hwnan
brain, seen from above at the midline. (Courtesy of Mark Williams, Dale Purves,
and Len White.)
The back cover reproduces the book's unit-opening images. See pages 2-t 188, 352,
476, 586, and IC-1.

NEUROSCIENCE, Fifth Edition

Copyright© 2012 by Sinauer Associates, Inc. All rights reserved.
This book may not be reproduced in whole or in part without permission.
Address inquiries and orders to
Sinauer Associates, Inc.
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Library of Congress Cataloging-in-Publication Data

Neuroscience I edited by Dale Purves ... Iet al.]. -- 5th ed.
p.cm.
Includes bibliographical references and index.
ISBN 978-0-87893-695-3 (casebound)
1. Neurosciences. I. Purves, Dale.
QP355.2.N487 2012
612.8--dc23 2011038400

Printed in U.SA

54321
CONTENTS IN BRIEF
1 Studying the Nervous System 1
Unit I Neural Signaling
2 Electrical Signals of Nerve Cells 25
3 Voltage-Dependent Membrane Permeability 41
4 Ion Channels and Transporters 57
5 Synaptic Transmission 77
6 Neurotransmitters and Their Receptors 109
7 Molecular Signaling within Neurons 141
8 Synaptic Plasticity 163
Unit II Sensation and Sensory Processing
9 The Somatic Sensory System: Touch and Proprioception 189
10 Pain 209
11 Vision: The Eye 229
12 Central Visual Pathways 257
13 The Auditory System 277
14 The Vestibular System 303
15 The Chemical Senses 321
Unit III Movement and Its Central Control
16 Lower Motor Neuron Circuits and Motor Control 353
17 Upper Motor Neuron Control of the Brainstem and Spinal Cord 375
18 Modulation of Movement by the Basal Ganglia 399
19 Modulation of Movement by the Cerebellum 417
20 Eye Movements and Sensory Motor Integration 435
21 The Visceral Motor System 451
Unit IV The Changing Brain
22 Early Brain Development 477
23 Construction of Neural Circuits 507
24 Modification of Neural Circuits as a Result of Experience 537
25 Repair and Regeneration in the Nervous System 559
Unit V Complex Brain Functions
26 Association Cortex and Cognition 587
27 Speech and Language 607
28 Sleep and Wakefulness 625
29 Emotions 647
30 Sex, Sexuality, and the Brain 669
31 Memory 695
Appendix: Survey of Human Neuroanatomy 717
Atlas: The Human Central Nervous System 745
CONTENTS

CHAPTER 1 Neural Circuits 10

The Organization of the Human Nervous System 13
Studying the Nervous System 1 Neural Systems 15
Overview 1 Structural Analysis of Neural Systems 15
Genetics, Genomics, and the Brain 1 Functional Analysis of Neural Systems 16
BOX 1A MODEL ORGANISMS IN NEUROSCIENCE 2 BOX 1B BRAIN IMAGING TECHNIQUES 18
The Cellular Components of the Nervous System 4 Analyzing Complex Behavior 21
Neurons 6 Summary 21
Glial Cells 7 Additional Reading 21
Cellular Diversity in the Nervous System 10

UNIT I
NEURAL SIGNALING 23
Summary 40
Additional Reading 40

CHAPTER 3
CHAPTER 2 Voltage-Dependent Membrane
Permeability 41
Electrical Signals of Nerve Cells 25
Overview 41
Overview 25 Ionic Currents across Nerve Cell Membranes 41
Electrical Signals of Nerve Cells 25
BOX 3A THE VOLTAGE CLAMP METHOD 42
Long-Distance Transmission of Electrical Signals 27
Two Types of Voltage-Dependent Ionic Currents 43
How Ion Movements Produce Electrical Signals 29
Two Voltage-Dependent Membrane Conductances 45
BOX 2A PASSIVE MEMBRANE PROPERTIES 30
Reconstruction of the Action Potential 47
Forces That Create Membrane Potentials 33
BOX 3B THRESHOLD 49
Electrochemical Equilibrium in an Environment with
More Than One Permeant Ion 34 Long-Distance Signaling by Means of Action
Potentials 51
The Ionic Basis of the Resting Membrane Potential 35
Increased Conduction Velocity as a Result of
BOX 2B THE REMARKABLE GIANT NERVE CELLS OF Myelination 51
SQUID 36 Summary 53
The Ionic Basis of Action Potentials 37
BOX 3C MULTIPLE SCLEROSIS 54
BOX 2C ACTION POTENTIAL FORM AND Additional Reading 55
NOMENCLATURE 39
VIII CONTENTS

CHAPTER 4 CHAPTER 6
Ion Channels and Transporters 57 Neurotransmitters and Their
Overview 57 Receptors 109
Ion Channels Underlying Action Potentials 57 Overview 109
BOX 4A THE PATCH CLAMP METHOD 59 Categories of Neurotransmitters 109
BOX 4B TOXINS THAT POISON ION CHANNELS 62 Acetylcholine 111
The Diversity of Ion Channels 63 BOX 6A NEUROTOXINS THAT ACT ON POSTSYNAPTIC
BOX 4C EXPRESSION OF ION CHANNELS IN XENOPUS RECEPTORS 114
OOCYTES 63 Glutamate 116
Voltage-Gated Ion Channels 64 BOX 6B MYASTHENIA GRAVIS: AN AUTOIMMUNE
Ligand-Gated Ion Channels 66 DISEASE OF NEUROMUSCULAR SYNAPSES 117
Stretch- and Heat-Activated Channels 66 BOX 6C EXCITOTOXICITY FOLLOWING ACUTE BRAIN
The Molecular Structure of Ion Channels 66 INJURY 121
Active Transporters Create and Maintain Ion GABA and Glycine 122
Gradients 69 BOX 6D EXCITATORY ACTIONS OF GABA IN THE
BOX 4D DISEASES CAUSED BY ALTERED ION DEVELOPING BRAIN 124
CHANNELS 70 The Biogenic Amines 125
Functional Properties of the Na+/K+ Pump 72 BOX 6E BIOGENIC AMINE NEUROTRANSMITTERS
The Molecular Structure of ATPase Pumps 72 AND PSYCHIATRIC DISORDERS 126
Summary 74 BOX 6F ADDICTION 128
Additional Reading 75 ATP and Other Purines 131
Peptide Neurotransmitters 132
CHAPTER 5 Unconventional Neurotransmitters 135
BOX 6G MARIJUANA AND THE BRAIN 137
Synaptic Transmission 77 Summary 139
Overview 77 Additional Reading 139
Electrical Synapses 78
Signal Transmission at Chemical Synapses 80 CHAPTER 7
Properties of Neurotransmitters 81
BOX 5A CRITERIA THAT DEFINE A
Molecular Signaling within
NEUROTRANSMITTER 84 Neurons 141
Quantal Release of Neurotransmitters 84 Overview 141
Release of Transmitters from Synaptic Vesicles 86 Strategies of Molecular Signaling 141
Local Recycling of Synaptic Vesicles 86 The Activation of Signaling Pathways 143
The Role of Calcium in Transmitter Secretion 88 Receptor Types 144
Molecular Mechanisms of Synaptic Vesicle Cycling 90 G-Proteins and Their Molecular Targets 145
BOX 5B DISEASES THAT AFFECT THE PRESYNAPTIC Second Messengers 147
TERMINAL 93 BOX 7A DYNAMIC IMAGING OF INTRACELLULAR
Neurotransmitter Receptors 96 SIGNALING 149
Postsynaptic Membrane Permeability Changes during Second Messenger Targets: Protein Kinases and
Synaptic Transmission 97 Phosphatases 151
Excitatory and Inhibitory Postsynaptic Potentials 101 Nuclear Signaling 153
Summation of Synaptic Potentials 102 BOX 7B DENDRITIC SPINES 154
BOX 5C THE “TRIPARTITE SYNAPSE” 104 Examples of Neuronal Signal Transduction 157
Summary 105 Summary 159
Additional Reading 106 Additional Reading 160
 CONTE NTS IX

CHAPTER 8 Mechanisms Underlying LTP 173

BOX 8B SILENT SYNAPSES 175
Synaptic Plasticity 163
Mechanisms Underlying LTD 177
Overview 163 Spike-Timing Dependent Plasticity 181
Short-Term Synaptic Plasticity 163
BOX 8C EPILEPSY: THE EFFECT OF PATHOLOGICAL
Long-Term Synaptic Plasticity Underlies Behavioral ACTIVITY ON NEURAL CIRCUITRY 182
Modification in Aplysia 166
Summary 183
Long-Term Potentiation at a Hippocampal Synapse 169
Additional Reading 184
BOX 8A GENETICS OF LEARNING AND MEMORY IN THE
FRUIT FLY 170

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 CHAPTER 10
The Somatic Sensory System: Pain 209
Touch and Proprioception 189 Overview 209
Overview 189 Nociceptors 209
Afferent Fibers Convey Somatic Sensory Information to Transduction and Transmission of Nociceptive
the Central Nervous System 189 Signals 211
Somatic Sensory Afferents Exhibit Distinct Functional BOX 10A CAPSAICIN 212
Properties 191
Central Pain Pathways Are Distinct from Mechano-
BOX 9A DERMATOMES 191 sensory Pathways 213
Mechanoreceptors Specialized to Receive Tactile BOX 10B REFERRED PAIN 214
Information 194
BOX 10C A DORSAL COLUMN PATHWAY FOR VISCERAL
Mechanoreceptors Specialized for Proprioception 196
PAIN 215
Central Pathways Conveying Tactile Information from
the Body: The Dorsal Column–Medial Lemniscal Parallel Pain Pathways 217
System 198 Pain and Temperature Pathways for the Face 218
Central Pathways Conveying Tactile Information from Other Modalities Mediated by the Anterolateral
the Face: The Trigeminothalamic System 200 System 220
Central Pathways Conveying Proprioceptive Information Sensitization 220
from the Body 200 Descending Control of Pain Perception 223
Central Pathways Conveying Proprioceptive Information BOX 10D PHANTOM LIMBS AND PHANTOM PAIN 222
from the Face 201
The Placebo Effect 223
The Somatic Sensory Components of the Thalamus 201
The Physiological Basis of Pain Modulation 224
Primary Somatic Sensory Cortex 202
Summary 226
BOX 9B PATTERNS OF ORGANIZATION WITHIN THE
Additional Reading 226
SENSORY CORTICES: BRAIN MODULES 203
X CONTENTS

CHAPTER 13
CHAPTER 11
The Auditory System 277
Vision: The Eye 229 Overview 277
Overview 229 Sound 277
Anatomy of the Eye 229 The Audible Spectrum 278
The Formation of Images on the Retina 230 A Synopsis of Auditory Function 279
BOX 11A MYOPIA AND OTHER REFRACTIVE ERRORS 231 BOX 13A FOUR CAUSES OF ACQUIRED
The Surface of the Retina 233 HEARING LOSS 279
Retinal Circuitry 233 BOX 13B MUSIC 280
BOX 11B THE BLIND SPOT 234 The External Ear 281
BOX 11C MACULAR DEGENERATION 235 The Middle Ear 282
BOX 11D RETINITIS PIGMENTOSA 237 BOX 13C SENSORINEURAL HEARING LOSS AND
Retinal Pigment Epithelium 238 COCHLEAR IMPLANTS 283
Phototransduction 238 The Inner Ear 284
Functional Specialization of the Rod and Cone Hair Cells and the Mechanoelectrical Transduction of
Systems 242 Sound Waves 287
Anatomical Distribution of Rods and Cones 244 The Ionic Basis of Mechanotransduction in Hair Cells 290
Cones and Color Vision 245 The Cochlear Amplifier 291
BOX 11E THE IMPORTANCE OF CONTEXT IN COLOR Tuning and Timing in the Auditory Nerve 291
PERCEPTION 246 BOX 13D THE SWEET SOUND OF DISTORTION 292
Retinal Circuits for Detecting Luminance Change 249 How Information from the Cochlea Reaches Targets in
BOX 11F THE PERCEPTION OF LIGHT INTENSITY 250
the Brainstem 293
Contribution of Retinal Circuits to Light Adaptation 253 Integrating Information from the Two Ears 294
Summary 255 Monaural Pathways from the Cochlear Nucleus to the
Nuclei of the Lateral Lemniscus 297
Additional Reading 256
Integration in the Inferior Colliculus 297
The Auditory Thalamus 297
CHAPTER 12 The Auditory Cortex 298
Central Visual Pathways 257 BOX 13E REPRESENTING COMPLEX SOUNDS IN THE
Overview 257 BRAINS OF BATS AND HUMANS 299
Central Projections of Retinal Ganglion Cells 257 Summary 301
The Retinotopic Representation of the Visual Field 259 Additional Reading 301
Visual Field Deficits 261
Spatiotemporal Tuning Properties of Neurons in Primary CHAPTER 14
Visual Cortex 263
The Vestibular System 303
Primary Visual Cortex Architecture 265
Overview 303
Combining Inputs from Two Eyes 267
The Vestibular Labyrinth 303
Division of Labor within the Primary Visual Pathway 269
BOX 14A A PRIMER ON TERMINOLOGY 304
BOX 12A RANDOM DOT STEREOGRAMS AND RELATED
AMUSEMENTS 270 Vestibular Hair Cells 305
The Functional Organization of Extrastriate Visual BOX 14B ADAPTATION AND TUNING OF VESTIBULAR
Areas 272 HAIR CELLS 306
Summary 274 The Otolith Organs: The Utricle and Saccule 307
Additional Reading 275 How Otolith Neurons Sense Tilts and Linear Accelera-
tions of the Head 309
The Semicircular Canals 310
How Semicircular Canal Neurons Sense Angular
Accelerations 312
 CONTE NTS XI

Central Pathways for Stabilizing Gaze, Head, and BOX 15A YOU ONLY GET ONE NOSE 326
Posture 312 Physiological and Behavioral Responses to Odorants 327
BOX 14C CLINICAL EVALUATION OF THE VESTIBULAR BOX 15B PHEROMONES, REPRODUCTION, AND THE
SYSTEM 314 VOMERONASAL SYSTEM 328
BOX 14D MAUTHNER CELLS IN FISH 316 Olfactory Epithelium and Olfactory Receptor
Vestibular Pathways to the Thalamus and Cortex 318 Neurons 329
Spatial Orientation Perception and Multisensory Odor Transduction and Odorant Receptor Proteins 332
Integration 318 BOX 15C THE “DOGTOR” IS IN 333
Summary 319 Physiological Mechanisms of Odor Transduction 334
Additional Reading 320 The Olfactory Bulb 336
Cortical Processing of Information Relayed from the
CHAPTER 15 Olfactory Bulb 340
The Organization of the Taste System 341
The Chemical Senses 321
Taste Perception in Humans 343
Overview 321
Taste Receptor Proteins and Transduction 345
The Organization of the Olfactory System 321
Neural Coding in the Taste System 347
Olfactory Perception in Humans 323
Trigeminal Chemoreception 349
Assessing Olfactory Function in the Laboratory or
Summary 349
Clinic 324
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
CHAPTER 16 STOMATOGASTRIC GANGLION 370
The Lower Motor Neuron Syndrome 372
Lower Motor Neuron Circuits and BOX 16D AMYOTROPHIC LATERAL SCLEROSIS 373
Motor Control 353 Summary 373
Overview 353 Additional Reading 374
Neural Centers Responsible for Movement 353
Motor Neuron–Muscle Relationships 355 CHAPTER 17
The Motor Unit 357
The Regulation of Muscle Force 358
Upper Motor Neuron Control of the
BOX 16A MOTOR UNIT PLASTICITY 359
Brainstem and Spinal Cord 375
The Spinal Cord Circuitry Underlying Muscle Stretch Overview 375
Reflexes 362 Organization of Descending Motor Control 375
The Influence of Sensory Activity on Motor Behavior 364 The Corticospinal and Corticobulbar Tracts 377
Other Sensory Feedback Affecting Motor BOX 17A PATTERNS OF FACIAL WEAKNESS AND
Performance 365 THEIR IMPORTANCE FOR LOCALIZING NEUROLOGICAL
Flexion Reflex Pathways 367 INJURY 379
Functional Organization of the Primary Motor Cortex 380
XII CONTENTS

BOX 17B WHAT DO MOTOR MAPS REPRESENT? 381 BOX 19B GENETIC ANALYSIS OF CEREBELLAR
BOX 17C SENSORY MOTOR TALENTS AND CORTICAL FUNCTION 431
SPACE 384 Summary 433
The Premotor Cortex 387 Additional Reading 433
Motor Control Centers in the Brainstem: Upper Motor
Neurons that Maintain Balance, Govern Posture, and CHAPTER 20
Orient Gaze 389
BOX 17D THE RETICULAR FORMATION 391
Eye Movements and Sensory Motor
Damage to Descending Motor Pathways: The Upper Integration 435
Motor Neuron Syndrome 395 Overview 435
BOX 17E MUSCLE TONE 397 What Eye Movements Accomplish 435
Summary 397 The Actions and Innervation of Extraocular Muscles 436
Additional Reading 398 BOX 20A THE PERCEPTION OF STABILIZED RETINAL
IMAGES 437
CHAPTER 18 Types of Eye Movements and Their Functions 438
Neural Control of Saccadic Eye Movements 440
Modulation of Movement by the Basal
BOX 20B SENSORY MOTOR INTEGRATION IN THE
Ganglia 399 SUPERIOR COLLICULUS 443
Overview 399 BOX 20C FROM PLACE CODES TO RATE CODES 445
Projections to the Basal Ganglia 399 Neural Control of Smooth Pursuit Movements 449
Projections from the Basal Ganglia to Other Brain Neural Control of Vergence Movements 449
Regions 402
Summary 450
Evidence from Studies of Eye Movements 404
Additional Reading 450
Circuits within the Basal Ganglia System 405
Dopamine Modulates Basal Ganglia Circuits 407
Hypokinetic and Hyperkinetic Movement Disorders 408
CHAPTER 21
BOX 18A PARKINSON’S DISEASE: AN OPPORTUNITY FOR The Visceral Motor System 451
NOVEL THERAPEUTIC APPROACHES 410 Overview 451
BOX 18B HUNTINGTON’S DISEASE 411 Early Studies of the Visceral Motor System 454
BOX 18C DEEP BRAIN STIMULATION 412 Distinctive Features of the Visceral Motor System 454
BOX 18D BASAL GANGLIA LOOPS AND NON-MOTOR The Sympathetic Division of the Visceral Motor
BRAIN FUNCTIONS 414 System 456
Summary 416 BOX 21A THE HYPOTHALAMUS 456
Additional Reading 416 The Parasympathetic Division of the Visceral Motor
System 460
CHAPTER 19 The Enteric Nervous System 461
Sensory Components of the Visceral Motor System 462
Modulation of Movement by the Central Control of Visceral Motor Functions 464
Cerebellum 417 BOX 21B HORNER’S SYNDROME 465
Overview 417 Neurotransmission in the Visceral Motor System 466
Organization of the Cerebellum 417 BOX 21C OBESITY AND THE BRAIN 466
Projections to the Cerebellum 419 Autonomic Regulation of Cardiovascular Function 468
Projections from the Cerebellum 421 Autonomic Regulation of the Bladder 470
Circuits within the Cerebellum 423 Autonomic Regulation of Sexual Function 472
Cerebellar Circuitry and the Coordination of Ongoing Summary 473
Movement 426
Additional Reading 474
BOX 19A PRION DISEASES 427
Further Consequences of Cerebellar Lesions 429
 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 Chemoattraction and Chemorepulsion 517
The Formation of Topographic Maps 519
Overview 477
Selective Synapse Formation 521
Formation of the Nervous System: Gastrulation and
Neurulation 477 Regulation of Neuronal Connections by Trophic
Interactions 523
BOX 22A STEM CELLS: PROMISE AND PERIL 479
Competitive Interactions and the Formation of Neuronal
Formation of the Major Brain Subdivisions 481 Connections 525
BOX 22B RHOMBOMERES 484 BOX 23C WHY DO NEURONS HAVE DENDRITES? 528
The Molecular Basis of Neural Induction 485 The Molecular Basis of Trophic Interactions 529
BOX 22C RETINOIC ACID: TERATOGEN AND INDUCTIVE BOX 23D THE DISCOVERY OF BDNF AND THE NEURO-
SIGNAL 488 TROPHIN FAMILY 530
Integrated Inductive Signals Establish Neuron Neurotrophin Signaling 533
Identity 489
Summary 535
The Initial Differentiation of Neurons and Glia 489
Additional Reading 535
BOX 22D NEUROGENESIS: WHERE, WHEN, AND
WHAT 492
Molecular Regulation of Neurogenesis 493 CHAPTER 24
The Generation of Neuronal Diversity 495 Modification of Neural Circuits as a
Molecular and Genetic Disruptions of Early Neural Result of Experience 537
Development 496
Overview 537
BOX 22E TRIPLE JEOPARDY: DISEASES ASSOCIATED WITH
Neural Activity and Brain Development 537
SONIC HEDGEHOG 497
Critical Periods 538
Neuronal Migration in the Peripheral Nervous
System 499 BOX 24A BUILT-IN BEHAVIORS 540
BOX 22F MIXING IT UP: LONG-DISTANCE NEURONAL BOX 24B BIRDSONG 541
MIGRATION 501 Cellular and Molecular Correlates of Activity-Dependent
Neuronal Migration in the Central Nervous System 502 Plasticity during Critical Periods 542
Molecular Mechanisms of Neuronal Migration and Critical Periods in Visual System Development 543
Cortical Migration Disorders 503 BOX 24C TRANSNEURONAL LABELING WITH RADIO-
Summary 503 ACTIVE AMINO ACIDS 544
Additional Reading 505 Effects of Visual Deprivation on Ocular Dominance 545
BOX 24D CORRELATION AS CAUSATION: LESSONS FROM
CHAPTER 23 A THREE-EYED FROG 549
Manipulating Competition 550
Construction of Neural Circuits 507
Amblyopia, Strabismus, and Critical Periods for Human
Overview 507 Vision 551
Neuronal Polarization: The First Step in Neural Circuit Evidence for Critical Periods in Other Sensory
Formation 507 Systems 552
XIV CONTENTS

The Development of Language: A Critical Period for a Regeneration of Peripheral Synapses 567
Distinctly Human Behavior 553 BOX 25A SPECIFIC REGENERATION OF SYNAPTIC
Human Brain Development, Activity-Dependent CONNECTIONS IN AUTONOMIC GANGLIA 568
Plasticity, and Critical Periods 554
Regeneration after Damage to the Central Nervous
Summary 556 System 570
Additional Reading 557 Cellular and Molecular Responses to Injury 570
BOX 25B CASUALTIES OF WAR AND SPORTS 571
CHAPTER 25 Axon Growth after Brain Injury 574
Repair and Regeneration in the Neurogenesis in the Mature Central Nervous System 575
Nervous System 559 Adult Neurogenesis in Non-Mammalian Vertebrates 576
Neurogenesis in the Adult Mammalian Brain 577
Overview 559
Cellular and Molecular Mechanisms of Adult
The Damaged Brain 559 Neurogenesis 578
Functional Reorganization without Repair 560 Adult Neurogenesis, Stem Cells, and Brain Repair in
Three Types of Neuronal Repair 561 Humans 580
Peripheral Nerve Regeneration 563 BOX 25C NUCLEAR WEAPONS AND NEUROGENESIS 581
The Cellular and Molecular Basis of Peripheral Nerve Summary 583
Repair 564
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
Speech and Language 607
Association Cortex and Cognition 587 Overview 607
Overview 587 Language Is Localized and Lateralized in the Brain 607
The Association Cortices 587 BOX 27A THE GENERATION OF SPEECH 608
An Overview of Cortical Structure 588 Aphasias 610
Unique Features of the Association Cortices 589
BOX 27B DO OTHER ANIMALS HAVE LANGUAGE? 611
BOX 26A CORTICAL LAMINATION 590
BOX 27C WORDS AND MEANING 613
The Parietal Association Cortex Mediates Attention 591
Confirmation of Language Lateralization and Other
“Attention Neurons” in the Monkey Parietal Cortex 594 Insights 615
The Temporal Association Cortex Mediates BOX 27D LANGUAGE AND HANDEDNESS 617
Recognition 596
The Search for Anatomical Differences between the Right
“Recognition Neurons” in the Monkey Temporal and Left Hemispheres 618
Cortex 598
Mapping Language Functions 619
The Frontal Association Cortex Mediates Planning and
Decision Making 599 The Role of the Right Hemisphere 622
BOX 26B NEUROPSYCHOLOGICAL TESTING 600 Sign Language 622
Summary 623
BOX 26C PSYCHOSURGERY 602
Additional Reading 623
 CONTE NTS XV

CHAPTER 30
CHAPTER 28
Sex, Sexuality, and the Brain 669
Sleep and Wakefulness 625 Overview 669
Overview 625 Sexual Dimorphisms and Sexually Dimorphic
Why Do Humans (and Many Other Animals) Sleep? 625 Behaviors 669
BOX 28A THE SLEEP STYLES OF DIFFERENT SPECIES 627 Sex, Gonads, Bodies, and Brains 671
The Circadian Cycle of Sleep and Wakefulness 628 BOX 30A THE SCIENCE OF LOVE (OR, LOVE IS A
BOX 28B MOLECULAR MECHANISMS OF BIOLOGICAL DRUG) 672
CLOCKS 629 Hormonal Influences on Sexual Dimorphism 674
Stages of Sleep 631 Primary Sexual Dimorphisms in the Brain 676
BOX 28C ELECTROENCEPHALOGRAPHY 632 Brain Dimorphisms and the Control of Reproductive
Behaviors 678
Physiological Changes in Sleep States 635
Structural and Functional Dimorphisms for Pregnancy
Other Possible Functions of Sleep and Dreaming 636
and Parenting 680
Neural Circuits Governing Sleep 637
BOX 30B THE GOOD MOTHER 683
BOX 28D CONSCIOUSNESS 638
Cellular and Molecular Basis of Sexually Dimorphic
Thalamocortical Interactions in Sleep 641 Structures and Behaviors 684
BOX 28E DRUGS AND SLEEP 641 Steroid Receptors and Responses in the Adult Brain 686
Sleep Disorders 643 Human Genetic Disorders of Genotypic and Phenotypic
BOX 28F CHRONIC FATIGUE SYNDROME 644 Sex 687
Summary 645 Sexual Orientation and the Brain: Molecular and Genetic
Analysis 688
Additional Reading 645
Sexual Orientation and Human Brain Structure 690
Sex-Based Differences in Cognitive Functions 691
CHAPTER 29 Summary 693
Emotions 647 Additional Reading 693
Overview 647
Physiological Changes Associated with Emotion 647 CHAPTER 31
BOX 29A FACIAL EXPRESSIONS: PYRAMIDAL AND EXTRA- Memory 695
PYRAMIDAL CONTRIBUTIONS 648
Overview 695
The Integration of Emotional Behavior 650
Qualitative Categories of Human Memory 695
The Limbic System 652
Temporal Categories of Memory 696
The Importance of the Amygdala 653
BOX 31A PHYLOGENETIC MEMORY 697
BOX 29B THE ANATOMY OF THE AMYGDALA 654
Memory Consolidation and Priming 698
BOX 29C THE REASONING BEHIND AN IMPORTANT
DISCOVERY 655
The Importance of Association in Information Storage 698
Conditioned Learning 699
BOX 29D FEAR AND THE HUMAN AMYGDALA:
A CASE STUDY 657 BOX 31B SAVANT SYNDROME 700
BOX 29E AFFECTIVE DISORDERS 659 Forgetting 702
The Relationship between Neocortex and Amygdala 660 Brain Systems Underlying Declarative Memory
Acquisition and Storage 703
Cortical Lateralization of Emotional Functions 661
BOX 31C CLINICAL CASES REVEAL AN ANATOMICAL SUB-
Emotion, Reason, and Social Behavior 662
STRATE FOR DECLARATIVE MEMORIES 703
Emotional Reinforcement and Addiction 663
Brain Systems Underlying Nondeclarative Memory
Summary 666 Acquisition and Storage 711
Additional Reading 667 Memory and Aging 712
BOX 31D ALZHEIMER’S DISEASE 713
Summary 714
Additional Reading 715
XVI CONTENTS

APPENDIX ATLAS
Survey of Human The Human Central Nervous
Neuroanatomy 717 System 745
Overview 717 Plate 1: Brain Surface 746
Neuroanatomical Plate 2: Coronal MR 748
Terminology 717 Plate 3: Axial MR 750
Basic Subdivisions of the Central Plate 4: Sagittal MR 754
Nervous System 718
Plate 5: Brainstem 756
External Anatomy of the Spinal
Cord 720 Plate 6: Spinal Cord 758
Internal Anatomy of the Spinal
Cord 721
Brainstem and Cranial Nerves 722 GLOSSARY G–1
Lateral Surface of the Brain 728
Dorsal and Ventral Surfaces of the Brain 729 ILLUSTRATION CREDITS IC–1
Midsagittal Surface of the Brain 730
INDEX I–1
Internal Anatomy of the Forebrain 731
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
CONTRIBUTORS

George J. Augustine, Ph.D.

David Fitzpatrick, Ph.D.
William C. Hall, Ph.D.
Anthony-Samuel LaMantia, Ph.D.
James 0. McNamara, M.D.
Richard D. Mooney, Ph.D.
Michael L. Platt, Ph.D.
Dale Purves, M.D.
Sidney A. Simon, Ph.D.
Leonard E. White, Ph.D.

UNIT EDITORS
UNIT I: George J. Augustine
UNIT II: David Fitzpatrick and Richard D. Mooney
UNIT III: Leonard E. White and William C. Hall
UNIT IV: Anthony-Samuel LaMantia
UNIT V: Dale Purves and Michael L. Platt