Why Do You Need this New Edition? If you’re wondering why you should buy this new edition of Physiology of Behavior, here are several good reasons: • Over 400 new research references. Biopsychology as a field evolves rapidly, with new research methods applied every year. The new research reported in this edition reflects the enormous advances made in research methods. Instructors will include this new material in your exams. • Updated illustrations. The author has revised existing art and prepared new art to illustrate research that is described for the first time in this edition. The result is a set of up-to-date, clear, consistent, and attractive illustrations. • NEW Review Questions are included at the end of each chapter so you can check your understanding of the chapter’s content. • Updated Section Summaries with Thought Questions. Summaries appear at the end of each major sec- tion so you have the chance to stop and review several times in each chapter. Section Summaries now include Thought Questions so you can test your understanding of the material. • NEW MyPsychLab combines original online learning applications with online assessments to help you engage in learning, assess your progress, and help you succeed. For each chapter of the text, MyPsychLab has a pre-test, post-test and chapter exam so you can get immediate feedback on your progress. You will receive a personalized study plan to help you succeed. MyPsychLab also contains an eText so you can access your textbook anytime, anywhere, including listening online. • NEW feature: Explore the Virtual Brain in MyPsychLab. This feature appears at the end of every chapter and directs you to relevant content in the Virtual Brain application in MyPsychLab. Virtual Brain is an interactive 3D application which allows you to take tours through different sections of the brain while using real life scenarios to explain behavior. This page intentionally left blank eleventh edition Physiology Behavior of Neil R. CaRlsoN University of Massachusetts, Amherst Boston Columbus Indianapolis New York San Francisco Upper Saddle River Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montreal Toronto Delhi Mexico City Sao Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo Editorial Director: Craig Campanella Editor in Chief: Jessica Mosher Acquisitions Editor: Amber Chow Senior Sponsoring Editor: Amber Mackey Editorial Assistant: Diane Szulecki Director of Marketing: Brandy Dawson Senior Marketing Manager: Nicole Kunzmann Senior Managing Editor: Maureen Richardson Project Manager: Annemarie Franklin Senior Operations Supervisor: Mary Fischer Operations Specialist: Diane Peirano Creative Director: Blair Brown Art Director Cover: Leslie Osher Cover Designer: Joseph DePinho Project Coordination, Text Design, and Electronic Page Makeup: Cenveo Publisher Services/Nesbitt Graphics Cover Art: iStock Photos Media Editor: Michael Halas Printer/Binder: Courier/Kendallville Cover Printer: Lehigh-Phoenix Color/Hagerstown Text Font: ITC New Baskerville Std. Copyright © 2013, 2010, 2007, 2004 by Pearson Education, Inc. All rights reserved. Printed in the United States of America. This publication is protected by Copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmis- sion in any form or by any means, electronic, mechanical, photocopying, record- ing, or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, One Lake Street, Upper Saddle River, New Jersey 07458 or you may fax your request to 201-236-3290. Library of Congress Cataloging-in-Publication Data available upon request. 10 9 8 7 6 5 4 3 2 1 Student Edition: ISBN-10: 0-205-23939-0 ISBN-13: 978-0-205-23939-9 Instructor’s Review Copy: ISBN-10: 0-205-23948-X ISBN-13: 978-0-205-23948-1 A la Carte: ISBN-10: 0-205-23981-1 ISBN-13: 978-0-205-23981-8 Brief Contents Chapter 1 Introduction 1 Chapter 2 Structure and Functions of Cells of the Nervous System 27 Chapter 3 Structure of the Nervous System 66 Chapter 4 Psychopharmachology 99 Chapter 5 Methods and Strategies of Research 130 Chapter 6 Vision 164 Chapter 7 Audition, the Body Senses, and the Chemical Senses 207 Chapter 8 Control of Movement 255 Chapter 9 Sleep and Biological Rhythms 288 Chapter 10 Reproductive Behavior 323 Chapter 11 Emotion 359 Chapter 12 Ingestive Behavior 393 Chapter 13 Learning and Memory 434 Chapter 14 Human Communication 479 Chapter 15 Neurological Disorders 516 Chapter 16 Schizophrenia and the Affective Disorders 552 Chapter 17 Anxiety Disorders, Autistic Disorder, Attention-Deficit/Hyperactivity Disorder, and Stress Disorders 584 Chapter 18 Drug Abuse 614 v This page intentionally left blank Contents 1 Introduction 1 Understanding Human Consciousness: Evolution of the Human Species 16 a Physiological approach 3 Evolution of Large Brains 19 Blindsight 4 ■ seCtioN sUmmaRy 21 Split Brains 4 ethical issues in Research with animals 22 Unilateral Neglect 7 Perception of Self 8 Careers in Neuroscience 26 ■ seCtioN sUmmaRy 8 ■ seCtioN sUmmaRy 24 the Nature of Behavioral Neuroscience 9 strategies for learning 25 The Goals of Research 10 Review Questions 26 Biological Roots of Behavioral Neuroscience 10 Explore the Virtual Brain in MyPsychLab 26 ■ seCtioN sUmmaRy 14 Natural selection and evolution 14 Functionalism and the Inheritance of Traits 14 Structure and Functions 2 of Cells of the Nervous System 27 Cells of the Nervous system 29 Activation of Receptors 56 Neurons 29 Postsynaptic Potentials 57 Supporting Cells 36 Termination of Postsynaptic Potentials 58 The Blood–Brain Barrier 39 Effects of Postsynaptic Potentials: Neural Integration 60 ■ seCtioN sUmmaRy 40 Autoreceptors 60 Communication Within a Neuron 41 Other Types of Synapses 61 Neural Communication: An Overview 41 Nonsynaptic Chemical Communication 62 Measuring Electrical Potentials of Axons 43 ■ seCtioN sUmmaRy 63 The Membrane Potential: Balance of Two Forces 45 Review Questions 65 The Action Potential 46 Explore the Virtual Brain in MyPsychLab 65 Conduction of the Action Potential 49 ■ seCtioN sUmmaRy 50 Communication Between Neurons 51 Structure of Synapses 52 Release of Neurotransmitter 54 vii viii Contents 3 Structure of the Nervous System 66 Basic Features of the Nervous system 67 the Peripheral Nervous system 93 An Overview 70 Spinal Nerves 93 Meninges 70 Cranial Nerves 93 The Ventricular System and Production The Autonomic Nervous System 94 of CSF 71 ■ seCtioN sUmmaRy 98 ■ seCtioN sUmmaRy 74 Review Questions 98 the Central Nervous system 74 Exploring the Virtual Brain in MyPsychLab 98 Development of the Central Nervous System 74 The Forebrain 80 The Midbrain 88 The Hindbrain 89 The Spinal Cord 91 ■ seCtioN sUmmaRy 92 4 Psychopharmacology 99 Principles of Psychopharmacology 101 Neurotransmitters and Neuromodulators 111 Pharmacokinetics 101 Acetylcholine 111 Drug Effectiveness 103 The Monoamines 114 Effects of Repeated Administration 104 Amino Acids 121 Placebo Effects 105 Peptides 123 ■ seCtioN sUmmaRy 106 Lipids 124 Nucleosides 125 sites of Drug action 106 Soluble Gases 126 Effects on Production of Neurotransmitters 107 ■ seCtioN sUmmaRy 127 Effects on Storage and Release of Neurotransmitters 108 Review Questions 129 Effects on Receptors 108 Explore the Virtual Brain in MyPsychLab 129 Effects on Reuptake or Destruction of Neurotransmitters 110 ■ seCtioN sUmmaRy 110 Contents ix 5 Methods and Strategies of Research 130 Experimental Ablation 132 neurochemical methods 156 Evaluating the Behavioral Effects Finding Neurons That Produce Particular of Brain Damage 132 Neurochemicals 156 Producing Brain Lesions 132 Localizing Particular Receptors 157 Stereotaxic Surgery 134 Measuring Chemicals Secreted in the Brain 158 Histological Methods 135 ■ SECtion SummAry 159 Tracing Neural Connections 139 Genetic methods 160 Studying the Structure of the Living Twin Studies 160 Human Brain 142 Adoption Studies 161 ■ SECtion SummAry 144 Genomic Studies 161 recording and Stimulating neural Activity 146 Targeted Mutations 161 Recording Neural Activity 146 Antisense Oligonucleotides 162 Recording the Brain’s Metabolic ■ SECtion SummAry 162 and Synaptic Activity 149 Stimulating Neural Activity 151 Review Questions 163 ■ SECtion SummAry 154 6 Vision 164 the Stimulus 166 Retinal Disparity 184 Color 184 Anatomy of the Visual System 167 Modular Organization of the Striate Cortex 185 The Eyes 167 Photoreceptors 169 ■ SECtion SummAry 186 Connections Between Eye and Brain 171 Analysis of Visual information: ■ SECtion SummAry 173 role of the Visual Association Cortex 187 Two Streams of Visual Analysis 187 Coding of Visual information in the retina 174 Perception of Color 190 Coding of Light and Dark 174 Perception of Form 191 Coding of Color 176 Perception of Movement 198 ■ SECtion SummAry 180 Perception of Spatial Location 201 Analysis of Visual information: ■ SECtion SummAry 204 role of the Striate Cortex 181 Review Questions 206 Anatomy of the Striate Cortex 181 Exploring the Virtual Brain in MyPsychLab 206 Orientation and Movement 181 Spatial Frequency 182