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GlobAl GlobAl edITIon edITIon eG dl ITo IobA nl P For these Global editions, the editorial team at Pearson has h y collaborated with educators across the world to address a wide range s i of subjects and requirements, equipping students with the best possible o l learning tools. This Global edition preserves the cutting-edge approach o g and pedagogy of the original, but also features alterations, customization, y and adaptation from the north American version. o f b e h a v Physiology of behavior i o r eT TWelFTH edITIon dW ITe Il Neil R. Carlson • Melissa A. Birkett oF T n H C a r l s o n • B i r k e t t This is a special edition of an established title widely used by colleges and universities throughout the world. Pearson published this exclusive edition for the benefit of students outside the United States and Canada. If you purchased this book within the United States or Canada, you should be aware that it has been imported without the approval of the Publisher or Author. Pearson Global Edition Carlson_12_1292158107_Final.indd 1 26/04/16 12:31 PM Physiology of Behavior twelfth edition global edition NEIL R. CARLSON University of Massachusetts, Amherst MELISSA A. BIRKETT Northern Arizona University Boston Columbus Indianapolis New York City San Francisco Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montréal Toronto Delhi Mexico City São Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo A01_CARL8105_12_GE_FM.indd 1 12/05/16 2:22 PM VP, Product Development: Dickson Musslewhite Director of Field Marketing: Jonathan Cottrell Senior Acquisitions Editor: Amber Chow Senior Product Marketing Manager: Editorial Assistant: Stephany Harrington Lindsey Prudhomme Gill Director, Content Strategy and Development: Brita Nordin Executive Field Marketing Manager: Kate Stewart Development Editor: Thomas Finn Marketing Assistant, Field Marketing: Amy Pfund Director, Project Management Services: Lisa Iarkowski Marketing Assistant, Product Marketing: Frank Alarcon Project Team Lead: Denise Forlow Operations Manager: Mary Fischer Project Manager: Shelly Kupperman Operations Specialist: Carol Melville Project Manager, Global Edition: Sudipto Roy Associate Director of Design: Blair Brown Senior Acquisitions Editor, Global Edition: Sandhya Ghoshal Interior Design: Kathryn Foot Senior Project Editor, Global Edition: Daniel Luiz Cover Design: Lumina Datamatics, Inc. Manager, Media Production, Global Edition: M. Vikram Kumar Cover Art: © allnow/Shutterstock Senior Manufacturing Controller, Production, Global Associate Digital Product Manager:: Christopher Fegan Edition: Trudy Kimber Digital Studio Project Manager: Pamela Weldin Program Team Lead: Amber Mackey Digital Studio Team Lead: Peggy Bliss Program Manager: Cecilia Turner Full-Service Project Management and Composition: Cenveo® Publisher Services 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 Edinburgh Gate Harlow Essex CM20 2JE England and Associated Companies throughout the world Visit us on the World Wide Web at: www.pearsonglobaleditions.com © 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. 10 9 8 7 6 5 4 3 2 1 14 13 12 11 10 Typeset in Palatino LT Pro by Cenveo® Publisher Services Printed and bound by Vivar in Malaysia. A01_CARL8105_12_GE_FM.indd 2 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 3 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 5 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 6 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 7 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 8 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 9 12/05/16 2:22 PM 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 A01_CARL8105_12_GE_FM.indd 10 12/05/16 2:22 PM