Download Neuroscience, Fifth Edition

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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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