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Physiology
of Behavior
twelfth edition
global edition
NEIL R. CARLSON
University of Massachusetts, Amherst
MELISSA A. BIRKETT
Northern Arizona University
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Acknowledgments of third party content appear on page 702, which constitutes an extension of this c opyright page.
Chapter Opener captions: Ch. 1: The human nervous system contains billions of neurons; Ch. 2: Neurons are the cells of the
nervous system that are specialized for communication; Ch. 3: The structures of the human nervous system are made up
of billions of neurons that make trillions of synapses; Ch. 4: Cross-section of the vagus nerve of the peripheral nervous
system; Ch. 5: Neurons in the cortex labeled with a fluorescent dye; Ch. 6: Cross-section of a retina. Photoreceptor cells are
visible at the top of the image; Ch. 7: Confocal microscopy image of neurons (green) and glia (red) in the vestibular pathway;
Ch 8: Cross-section of the cerebellum; Ch. 9: Cross-section of the hypothalamus of a mouse; Ch. 10: Cross-section of the
pituitary gland (left) attached to the hypothalamus (right); Ch. 11: Example of pyramidal neurons found in the hippocampus;
Ch. 12: Color-enhanced transmission electron micrograph of portions of two adipose cells and associated connective tissue
in a rat; Ch. 13: New neurons in the mouse hippocampus are labeled with green fluorescence; Ch. 14: Scanning electron
microscope image of a neuron in the cortex; Ch. 15: Neurons derived from mouse embryonic stem cells. Tyrosine hydroxylase
(TH, a dopamine-synthesizing enzyme) is labeled in red; TH-containing neurons degenerate in Parkinson’s disease; green
labels a protein that’s found in all neurons; blue labels the nuclei of all cells; Ch. 16: Neurons in the mouse hippocampus;
Ch. 17: Cross-section of the adrenal medulla; Ch. 18: Neurons in the CA1 region of the hippocampus from a transgenic
mouse stained for the CB1 cannabinoid receptor (red) and cell nuclei (blue).
Pearson Education Limited
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© Pearson Education Limited 2017
The rights of Neil R. Carlson and Melissa A. Birkett to be identified as the authors of this work have been asserted by them
in accordance with the Copyright, Designs and Patents Act 1988.
Authorized adaptation from the United States edition, entitled Physiology of Behavior, 12th edition, ISBN 978-0-13-408091-8, by Neil R.
Carlson and Melissa A. Birkett, published by Pearson Education © 2017.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form
or by any means, electronic, mechanical, photocopying, recording or otherwise, without either the prior written permission
of the publisher or a license permitting restricted copying in the United Kingdom issued by the Copyright Licensing Agency
Ltd, Saffron House, 6–10 Kirby Street, London EC1N 8TS.
All trademarks used herein are the property of their respective owners. The use of any trademark in this text does not vest in
the author or publisher any trademark ownership rights in such trademarks, nor does the use of such trademarks imply any
affiliation with or endorsement of this book by such owners.
ISBN 10: 1-292-15810-7
ISBN 13: 978-1-292-15810-5
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library.
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Typeset in Palatino LT Pro by Cenveo® Publisher Services
Printed and bound by Vivar in Malaysia.
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Brief Contents
1 Introduction 15
2 Structure and Functions of Cells of the Nervous System 35
3 Structure of the Nervous System 70
4 Psychopharmacology 102
5 Methods and Strategies of Research 132
6 Vision 163
7 Audition, the Body Senses, and the Chemical Senses 202
8 Control of Movement 245
9 Sleep and Biological Rhythms 275
10 Reproductive Behavior 310
11 Emotion 344
12 Ingestive Behavior 380
13 Learning and Memory 419
14 Human Communication 460
15 Neurological Disorders 495
16 Schizophrenia and the Affective Disorders 533
17 Stress, Anxiety, and Neurodevelopmental Disorders 566
18 Substance Abuse 602
3
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Contents
Preface 11 Activation of Receptors 61
Postsynaptic Potentials 62
Termination of Postsynaptic Potentials 63
Effects of Postsynaptic Potentials: Neural Integration 64
Autoreceptors 66
Other Types of Synapses 66
Other Forms of Chemical Communication 67
1 Introduction 15
Foundations of Behavioral Neuroscience 17
The Goals of Research 18
Biological Roots of Behavioral Neuroscience 18
Natural Selection and Evolution 23
Functionalism and the Inheritance of Traits 23 3 Structure of the Nervous System 70
Evolution of Large Brains 25
Ethical Issues in Research with Humans Basic Features of the Nervous System 72
and Other Animals 28 Anatomical Directions 73
Research with Animals 28 Meninges 75
Research with Humans 29 The Ventricular System and Production of CSF 75
The Future of Neuroscience: Development of the Nervous System 78
Careers and Strategies for Learning 31 An Overview of Brain Development 78
Careers in Neuroscience 31 Prenatal Brain Development 78
Strategies for Learning 31 Postnatal Brain Development 82
Structure and Function of the Central Nervous System 84
The Forebrain 84
The Midbrain 92
The Hindbrain 93
The Spinal Cord 94
Structure and Function of the Peripheral Nervous System 96
Cranial Nerves 96
2 Structure and Functions of Spinal Nerves 97
The Autonomic Nervous System 98
Cells of the Nervous System 35
Cells of the Nervous System 37
The Nervous System: An Overview 37
Neurons 38
Supporting Cells 43
The Blood–Brain Barrier 46
Communication Within a Neuron 47
Neural Communication: An Overview 47
4 Psychopharmacology 102
Measuring Electrical Potentials of Axons 49
The Membrane Potential 50 Principles of Psychopharmacology 104
The Action Potential 52 An Overview of Psychopharmacology 104
Conduction of the Action Potential 55 Pharmacokinetics 105
Communication Between Neurons 58 Drug Effectiveness 107
Structure of Synapses 58 Effects of Repeated Administration 108
Release of Neurotransmitters 59 Placebo Effects 109
5
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6 Contents
Sites of Drug Action 110 Anatomy of the Eye 167
Effects on Production of Neurotransmitters 111 Photoreceptors 168
Effects on Storage and Release of Neurotransmitters 111 Transduction 169
Effects on Receptors 112 Central and Peripheral Vision 171
Effects on Reuptake or Destruction of Neurotransmitters 113 The Optic Nerves 172
Overview of the Visual Pathway 173
Neurotransmitters and Neuromodulators 114
Amino Acids 115 Brain Regions Involved in Visual Processing 175
Acetylcholine 117 Lateral Geniculate Nucleus 175
The Monoamines 120 Striate Cortex 175
Extrastriate Cortex 177
Peptides 127
Lipids 128 Perception of Color 179
Role of the Retinal Ganglion Cells in
Light/Dark Perception 179
Role of the Retina in Color Perception 181
Role of the Striate Cortex 184
Role of the Extrastriate Cortex 184
Perception of Form 187
Role of the Striate Cortex 187
Role of the Extrastriate Cortex 189
5 Methods and Strategies of Research 132 Perception of Spatial Location 194
Role of the Retina 194
Experimental Ablation 135
Role of the Striate Cortex 195
Evaluating the Behavioral Effects of Brain Damage 135
Role of the Extrastriate Cortex 195
Producing Brain Lesions 135
Perception of Orientation and Movement 197
Stereotaxic Surgery 136
Role of the Striate Cortex 197
Histological Methods 138 Role of the Extrastriate Cortex 197
Tracing Neural Connections 140
Studying the Structure of the Living Human Brain 144
Recording and Stimulating Neural Activity 147
Recording Neural Activity 147
Recording the Brain’s Metabolic and Synaptic Activity 150
Stimulating Neural Activity 152
Neurochemical Methods 155
Finding Neurons That Produce
7 Audition, the Body Senses,
Particular Neurochemicals 155
Localizing Particular Receptors 157 and the Chemical Senses 202
Measuring Chemicals Secreted in the Brain 157
Audition 204
Genetic Methods 159
The Stimulus 204
Twin Studies 160
Anatomy of the Ear 205
Adoption Studies 160
Auditory Hair Cells and the Transduction
Genomic Studies 160 of Auditory Information 207
Targeted Mutations 160 The Auditory Pathway 208
Antisense Oligonucleotides 161 Perception of Pitch 211
Perception of Loudness 212
Perception of Timbre 212
Perception of Spatial Location 213
Perception of Complex Sounds 216
Perception of Music 217
Vestibular System 220
Anatomy of the Vestibular Apparatus 221
The Vestibular Pathway 222
6 Vision 163
Somatosenses 223
The Eye 166 The Stimuli 223
Introduction to Sensation and Perception 166 Anatomy of the Skin and Its Receptive Organs 224
The Stimulus: Light 166 Perception of Cutaneous Stimulation 225
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Contents 7
The Somatosensory Pathways 227 Why Do We Sleep? 282
Perception of Pain 229 Functions of Slow-Wave Sleep 283
Gustation 235 Functions of REM Sleep 285
The Stimuli 235 Sleep and Learning 285
Anatomy of the Taste Buds and Gustatory Cells 236 Physiological Mechanisms of Sleep and Waking 288
Perception of Gustatory Information 236 Neural Control of Sleep 288
The Gustatory Pathway 238 Neural Control of Arousal 289
Olfaction 239 Neural Control of Sleep/Wake Transitions 293
The Stimulus and Anatomy of the Olfactory Apparatus 240 Neural Control of Transition to REM 295
Transduction of Olfactory Information 241 Disorders of Sleep 298
Perception of Specific Odors 242 Insomnia 298
Narcolepsy 299
REM Sleep Behavior Disorder 301
Problems Associated with Slow-Wave Sleep 301
Biological Clocks 303
Circadian Rhythms and Zeitgebers 303
The Suprachiasmatic Nucleus 304
Control of Seasonal Rhythms:
The Pineal Gland and Melatonin 307
8 Control of Movement 245
Changes in Circadian Rhythms: Shift Work and Jet Lag 308
Skeletal Muscle 247
Anatomy 247
The Physical Basis of Muscular Contraction 249
Sensory Feedback from Muscles 250
Control of Movement by the Spinal Cord 252
The Monosynaptic Stretch Reflex 252
The Gamma Motor System 252
Polysynaptic Reflexes 254
10 Reproductive Behavior 310
Control of Movement by the Brain 255
Cortical Structures 255 Sexual Development 312
Cortical Control of Movement: Descending Pathways 257 Production of Gametes and Fertilization 312
Planning and Initiating Movements: Role Development of the Sex Organs 313
of the Motor Association Cortex 259
Sexual Maturation 315
Subcortical Structures 263
Hormonal Control of Sexual Behavior 318
Complex Motor Behavior 269
Hormonal Control of Female Reproductive Cycles 318
Imitating and Comprehending Movements: Role
Hormonal Control of Sexual Behavior of
of the Mirror Neuron System 269
Laboratory Animals 319
Control of Reaching and Grasping: Role
Organizational Effects of Androgens on Behavior:
of the Parietal Cortex 271
Masculinization and Defeminization 321
Deficits of Skilled Movements: The Apraxias 273
Human Sexual Behavior 321
Limb Apraxia 273
Effects of Pheromones 323
Constructional Apraxia 273
Neural Control of Sexual Behavior 327
Males 327
Females 330
Formation of Pair Bonds 330
Sexual Orientation 332
Activational and Organizational Effects of Hormones 333
Role of Androgens 333
Cloacal Exstrophy 334
The Sexually Dimorphic Brain 334
9 Sleep and Biological Rhythms 275
Role of Prenatal Environment in Sexual Orientation 336
What Is Sleep? 277 Heredity and Sexual Orientation 336
Stages of Sleep 278 Parental Behavior 338
Brain Activity During Sleep 280 Maternal Behavior of Rodents 338
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8 Contents
Hormonal Control of Maternal Behavior 339 The Long-Term Reservoir 389
Neural Control of Maternal Behavior 339 Fasting Phase 389
Neural Control of Paternal Behavior 342 Absorptive Phase 389
Eating: Signals to Start a Meal 391
Signals from the Digestive System 391
Metabolic Signals 392
Eating: Signals to Stop a Meal 394
Short-Term Satiety 395
Signals from Environmental Factors 396
Signals from Sensory Factors 396
Signals from Gastric Factors 396
11 Emotion 344 Signals from Intestinal Factors 396
Signals from Liver Factors 397
Fear 346
Signals from Insulin 398
Components of Emotional Response 346
Long-Term Satiety: Signals from
Research with Laboratory Animals 347 Adipose Tissue 398
Research with Humans 351 Brain Mechanisms 400
Aggression 353 Brain Stem 400
Research with Laboratory Animals 353 Hypothalamus 400
Research with Humans 354 Obesity 406
Hormonal Control of Aggressive Behavior 355 Possible Causes 407
Impulse Control 360 Treatment 409
Role of the vmPFC 360 Eating Disorders 413
Brain Development and Impulse Control 361 Possible Causes 415
Crime and Impulse Control 362 Treatment 417
Serotonin and Impulse Control 362
Moral Decision Making 363
Communication of Emotions 365
Facial Expression of Emotions:
Innate Responses 365
Neural Basis of the Communication
of Emotions: Recognition 366
Neural Basis of the Communication
of Emotions: Expression 372 13 Learning and Memory 419
Feelings of Emotions 375
Overview of Learning and Memory 422
The James-Lange Theory 376
Types of Learning 422
Feedback from Emotional Expressions 377
Types of Memory 424
Stimulus-Response Learning 427
Classical Conditioning 427
Operant Conditioning 429
Motor Learning 434
Role of the Cortex 434
Role of the Basal Ganglia 434
Perceptual Learning 435
12 Ingestive Behavior 380 Role of the Cortex 435
Retaining Perceptual Information
Drinking 382 in Short-Term Memory 436
Physiological Regulatory Mechanisms 382 Relational Learning 438
Two Types of Thirst 383 Role of the Hippocampus 438
Neural Mechanisms of Thirst 386 Role of the Cortex 443
Eating: What Is Metabolism? 388 Amnesia 444
The Short-Term Reservoir 388 Role of the Hippocampus 444
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Contents 9
Stimulus-Response Learning 446 Cerebrovascular Accidents 503
Motor Learning 446 Causes 503
Perceptual Learning 447 Treatments 504
Relational Learning 448 Traumatic Brain Injury 508
Long-Term Potentiation 452 Causes 508
Induction of Long-Term Potentiation 453 Treatments 509
Role of NMDA Receptors 453 Disorders of Development 510
Role of AMPA Receptors 456 Toxic Chemicals 510
Role of Synaptic Changes 457 Inherited Metabolic Disorders 510
Down Syndrome 512
Degenerative Disorders 514
Transmissible Spongiform
Encephalopathies 514
Parkinson’s Disease 515
Huntington’s Disease 520
Amyotrophic Lateral Sclerosis 522
Multiple Sclerosis 522
14 Human Communication 460
Dementia 524
Korsakoff’s Syndrome 528
Language Production and Comprehension:
Disorders Caused by Infectious
Brain Mechanisms 462
Diseases 530
Lateralization 463
Encephalitis 530
Language Production 464
Meningitis 531
Language Comprehension 465
Bilingualism 466
Prosody 467
Recognition of People’s Voices 467
Disorders of Language Production
and Comprehension 469
Disorders of Language Production:
Broca’s Aphasia 469
Disorders of Language Comprehension:
Wernicke’s Aphasia 472 16 Schizophrenia and the Affective
Conduction Aphasia 477
Disorders 533
Aphasia in People Who Are Deaf 479
Stuttering 480 Schizophrenia 535
Disorders of Reading and Writing 483 Description 535
Relation to Aphasia 483 Heritability 537
Pure Alexia 483 Environmental Factors 540
Toward an Understanding of Reading 485 Anomalies in Schizophrenia 542
Toward an Understanding of Writing 491 The Mesolimbic Dopamine
Pathway: Positive Symptoms 545
The Mesocortical Dopamine Pathway:
Negative and Cognitive Symptoms 547
Affective Disorders 551
Description 551
Heritability 552
Biological Treatments 552
Role of the Frontal Cortex 557
15 Neurological Disorders 495 The Monoamine Hypothesis 558
Role of the 5-HT Transporter 559
Tumors and Seizures 497
Role of Neurogenesis 560
Tumors 497
Role of Circadian
Seizures 500 Rhythms 560
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10 Contents
17 Stress, Anxiety, and 18 Substance Abuse 602
Neurodevelopmental Disorders 566
Common Features of Substance Abuse 605
Stress 568 Positive Reinforcement 606
Physiology of the Stress Response 569 Negative Reinforcement 611
Health Effects of Long-Term Stress 570 Heredity 615
Effects of Stress on the Brain 571 Alcohol 616
Psychoneuroimmunology 574 Nicotine 616
Posttraumatic Stress Disorder 578 Stimulants 616
Symptoms 578 Commonly Abused Drugs 617
Heritability 578 Opiates 617
Brain Changes 579 Stimulants 619
Treatment 581 Nicotine 621
Anxiety Disorders 582 Alcohol 624
Symptoms 582 Cannabis 625
Heritability 584 Treatment for Substance Abuse 628
Brain Changes 584 Opiates 629
Treatment 584 Stimulants 629
Obsessive-Compulsive Disorder 587 Nicotine 630
Symptoms 587 Alcohol 631
Heritability 588 Brain Stimulation 631
Brain Changes 589
Treatment 589
Autism Spectrum Disorder 592
Symptoms 593
Glossary 633
Heritability 593
Brain Changes 594 References 651
Attention-Deficit/Hyperactivity Disorder 597 Credits 702
Symptoms 597
Name Index 709
Heritability 598
Brain Changes 599 Subject Index 723
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