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Pinel, John P. J.
Biopsychology / John P.J. Pinel, University of British Columbia.—Ninth edition.
pages cm
Includes bibliographical references and index.
ISBN 978-0-205-91557-6 (alk. paper)
1. Psychobiology—Textbooks. I. Title.
QP360.P463 2015
612.8—dc23
2013030685
10 9 8 7 6 5 4 3 2 1
Part Four
Brain Plasticity
9 Development of the Nervous System 213
From Fertilized Egg to You
iii
Preface xvi
To the Student xxiii
About the Author xxiii
Part Two
The Case of Jimmie G.,
the Man Frozen in Time 3 Foundations of Biopsychology
Four Major Themes of This Text 3
1.1 What Is Biopsychology? 4 2 Evolution,
Genetics, and
1.2 What Is the Relation between Experience 20
Biopsychology and the Other Thinking about the
Disciplines of Neuroscience? 4 Biology of Behavior
1.3 What Types of Research Characterize
2.1 Thinking about the Biology of Behavior:
the Biopsychological Approach? 4
From Dichotomies to Interactions 21
Human and Nonhuman Subjects 5
Is It Physiological, or Is It Psychological? 21
Experiments and Nonexperiments 5
Is It Inherited, or Is It Learned? 22
Pure and Applied Research 7
Problems with Thinking about the
1.4 What Are the Divisions of Biopsychology? 8 Biology of Behavior in Terms of Traditional
Physiological Psychology 8 Dichotomies 22
Psychopharmacology 9 The Case of the Man Who Fell Out of Bed 23
Neuropsychology 9 Case of the Chimps with Mirrors 23
The Case of Mr. R., the Brain-Damaged The Case of the Thinking Student 24
Student Who Switched to Architecture 9 2.2 Human Evolution 24
Psychophysiology 9 Evolution and Behavior 26
Cognitive Neuroscience 10 Course of Human Evolution 27
Comparative Psychology 11 Thinking about Human Evolution 30
1.5 Converging Operations: How Do Evolution of the Human Brain 32
Biopsychologists Work Together? 12 Evolutionary Psychology: Understanding
Mate Bonding 33
1.6 Scientific Inference: How Do
Biopsychologists Study the Thinking about Evolutionary Psychology 34
Unobservable Workings of the Brain? 13 2.3 Fundamental Genetics 35
1.7 Critical Thinking about Mendelian Genetics 35
Biopsychological Claims 14 Chromosomes: Reproduction and
Case 1: José and the Bull 15 Recombination 36
iv
3 Anatomy of the
Nervous System 51 4.4 Conduction of Action Potentials
Ionic Basis of Action Potentials
82
82
Systems, Structures,
and Cells That Make Up Refractory Periods 83
Your Nervous System Axonal Conduction of Action Potentials 83
Conduction in Myelinated Axons 83
3.1 General Layout of the Nervous System 52 The Velocity of Axonal Conduction 84
Divisions of the Nervous System 52 Conduction in Neurons without Axons 84
Meninges, Ventricles, and Cerebrospinal The Hodgkin-Huxley Model in
Fluid 53 Perspective 85
Blood–Brain Barrier 54
4.5 Synaptic Transmission: Chemical
3.2 Cells of the Nervous System 56 Transmission of Signals among
Anatomy of Neurons 56 Neurons 85
Glia: The Forgotten Cells 56 Structure of Synapses 85
Synthesis, Packaging, and Transport
3.3 Neuroanatomical Techniques of Neurotransmitter Molecules 86
and Directions 61 Release of Neurotransmitter Molecules 87
Neuroanatomical Techniques 61 Activation of Receptors by
Directions in the Vertebrate Nervous Neurotransmitter Molecules 87
System 62 Reuptake, Enzymatic Degradation, and
3.4 Spinal Cord 64 Recycling 89
Glia, Gap Junctions, and Synaptic
3.5 Five Major Divisions of the Brain 64 Transmission 90
5 The Research
Methods
Tests of the Common Neuropsychological
Test Battery 118
of Biopsychology 100 Tests of Specific Neuropsychological
Understanding What Function 119
Biopsychologists Do Frontal-Lobe Function 120
6
Eye Movement 108 The Visual System 129
Skin Conductance 109 How We See
Cardiovascular Activity 109 The Case of Mrs.
5.3 Invasive Physiological Research Richards: Fortification
Methods 109 Illusions and the
Astronomer 130
Stereotaxic Surgery 110
Lesion Methods 110 6.1 Light Enters the Eye and Reaches
Electrical Stimulation 111 the Retina 131
Invasive Electrophysiological Recording Pupil and the Lens 132
Methods 111 Eye Position and Binocular Disparity 133
7.4 Chemical Senses: Smell and Taste 179 Ventromedial Corticospinal Tract and
Olfactory System 180 Ventromedial Cortico-Brainstem-Spinal
Gustatory System 181 Tract 199
Brain Damage and the Chemical Senses 182 Comparison of the Two Dorsolateral
Motor Pathways and the Two Ventromedial
Broad Tuning vs. Narrow Tuning 182 Motor Pathways 200
7.5 Selective Attention 184
8.7 Sensorimotor Spinal Circuits 202
Change Blindness 184
Muscles 202
Neural Mechanisms of Attention 185
Receptor Organs of Tendons
Simultanagnosia 186 and Muscles 203
Themes Revisited 186 Stretch Reflex 204
Think about It 186 Withdrawal Reflex 205
Key Terms 187 Reciprocal Innervation 205
Quick Review 187 Recurrent Collateral Inhibition 205
Walking: A Complex Sensorimotor
Reflex 206
8 The Sensorimotor
System 188
8.8 Central Sensorimotor Programs
and Learning 207
How You Move
Central Sensorimotor Programs
The Case of Rhonda, the Are Capable of Motor Equivalence 208
Dexterous
Sensory Information That Controls Central
Cashier 189
Sensorimotor Programs Is Not Necessarily
8.1 Three Principles of Sensorimotor Conscious 208
Function 190 Central Sensorimotor Programs
The Sensorimotor System Is Hierarchically Can Develop without Practice 209
Organized 190 Practice Can Create Central Sensorimotor
Motor Output Is Guided by Sensory Input 190 Programs 209
The Case of G.O., the Man with Too Little Functional Brain Imaging of Sensorimotor
Feedback 190 Learning 209
Learning Changes the Nature and Locus The Case of Rhonda, Revisited 211
of Sensorimotor Control 191 Themes Revisited 211
General Model of Sensorimotor System Think about It 211
Function 191
Key Terms 211
8.2 Sensorimotor Association Cortex 191 Quick Review 212
Posterior Parietal Association Cortex 191
The Case of Mrs. S., the Woman Who
Turned in Circles 192 Part Four
Dorsolateral Prefrontal Association Cortex 193
Brain Plasticity
8.3 Secondary Motor Cortex 194
Identifying the Areas of Secondary
Motor Cortex 194 9 Development of the
Nervous System 213
Mirror Neurons 195 From Fertilized
Egg to You
8.4 Primary Motor Cortex 196
The Case of Genie 214
Belle: The Monkey That Controlled
a Robot with Her Mind 197
9.1 Phases of Neurodevelopment 214
8.5 Cerebellum and Basal Ganglia 198
Induction of the Neural Plate 215
Cerebellum 198
Neural Proliferation 215
Basal Ganglia 198
Migration and Aggregation 216
8.6 Descending Motor Pathways 199 Axon Growth and Synapse Formation 217
Dorsolateral Corticospinal Tract and Neuron Death and Synapse
Dorsolateral Corticorubrospinal Tract 199 Rearrangement 220
10
Brain Damage and Key Terms 258
Neuroplasticity 233 Quick Review 258
Can the Brain
Recover from Damage?
The Ironic Case of 11 Learning,
Memory,
Professor P. 234 and Amnesia 259
How Your Brain Stores
10.1 Causes of Brain Damage 235 Information
Brain Tumors 235
Cerebrovascular Disorders: Strokes 235
11.1 Amnesic Effects of Bilateral Medial
Closed-Head Injuries 237
Temporal Lobectomy 260
The Case of Jerry Quarry, Ex-Boxer 238
The Case of H.M., the Man Who
Infections of the Brain 238 Changed the Study of Memory 260
Neurotoxins 239 Formal Assessment of H.M.’s Anterograde
Genetic Factors 239 Amnesia: Discovery of Unconscious
Programmed Cell Death 239 Memories 261
13 Hormones
and Sex 314
Female Reproduction-Related Behavior
and Gonadal Hormones 331
What’s Wrong with the Anabolic Steroid Abuse 333
Mamawawa?
13.6 Brain Mechanisms of Sexual Behavior 334
Men-Are-Men-and-
Women-Are-Women Cortex and Sexual Activity 334
Assumption 315 Hypothalamus and Sexual Activity 334
Developmental and Activational Effects Amygdala and Sexual Activity 335
of Sex Hormones 315 Ventral Striatum and Sexual Activity 336
13.1 The Neuroendocrine System 315 13.7 Sexual Orientation and Sexual Identity 336
Glands 316 Sexual Orientation and Genes 336
Gonads 316 Sexual Orientation and Early Hormones 336
16 Lateralization,
Language, and
The Wernicke-Geschwind Model
16.6 Wernicke-Geschwind Model:
411
17 Biopsychology of
Emotion, Stress,
18 Biopsychology of
Psychiatric
Disorders 446
and Health 423 The Brain Unhinged
Fear, the Dark Side
of Emotion
W
elcome to the Ninth Edition of Biopsychology! Effective Use of Case Studies Biopsychology fea-
This edition builds on the strengths of its pre- tures many carefully selected case studies, which are
decessors, but it also takes important new steps: highlighted in the text. These provocative cases stimulate
In addition to covering many new cutting-edge research interest, promote retention, and allow students to learn
topics, it sharpens its focus on the human element of how biopsychological principles apply to the diagnosis
biopsychology and on promoting student thinking. Most and treatment of brain disorders.
importantly, this is the first edition of Biopsychology to
focus on epigenetics. It introduces this new field, summa- Remarkable Illustrations The illustrations in Bio
rizes current knowledge, and most importantly stresses psychology are special. Each one was conceptualized and
the relevance of epigenetic concepts to issues of human meticulously designed to clarify and reinforce the text by
brain and behavior. a uniquely qualified scientist–artist team: Pinel and his
The Ninth Edition of Biopsychology is a clear, engag- artist/designer wife, Maggie Edwards.
ing introduction to current biopsychological theory and Focus on Behavior In some biopsychological text-
research. It is intended for use as a primary text in one- books, the coverage of neurophysiology, neurochemistry,
or two-semester courses in biopsychology—variously ti- and neuroanatomy subverts the coverage of behavioral
tled Biopsychology, Physiological Psychology, Brain and research. Biopsychology gives top billing to behavior: It
Behavior, Psychobiology, Behavioral Neuroscience, or stresses that neuroscience is a team effort and that the
Behavioral Neurobiology. unique contribution made by biopsychologists to this
The defining feature of Biopsychology is its unique effort is their behavioral expertise.
combination of biopsychological science and personal,
reader-oriented discourse. It is a textbook that is “un- Emphasis on the Scientific Method Biopsychology
textbooklike.” Instead of presenting the concepts of bio- emphasizes the scientific method. It portrays the scientific
psychology in the usual textbook fashion, it addresses method as a means of answering questions that is as appli-
students directly and interweaves the fundamentals of cable in daily life as in the laboratory. And Biopsychology
the field with clinical case studies, social issues, per- emphasizes that being a scientist is fun.
sonal implications, useful metaphors, and memorable
anecdotes. Discussion of Personal and Social Implications
Several chapters of Biopsychology—particularly those on
eating, sleeping, sex, and drug addiction—carry strong
Key Features Maintained personal and social messages. In these chapters, students
in the Ninth Edition are encouraged to consider the relevance of biopsycho-
logical research to their lives outside the classroom.
The following are features that have characterized recent
editions of Biopsychology and have been maintained or Engaging, Inspiring Voice Arguably the strongest
expanded in this edition. pedagogical feature of Biopsychology is its personal tone.
Pinel addresses students directly and talks to them with
Emphasis on Broad Themes The emphasis of Bio warmth, enthusiasm, and good humor about recent ad-
psychology is “the big picture.” Four broad themes are vances in biopsychological science. Many students report
highlighted throughout the text by distinctive tabs: being engaged and inspired by this approach.
(1) thinking creatively, (2) clinical implications, (3) evo-
lutionary perspective, and (4) neuroplasticity. A Themes
Revisited section at the end of each chapter briefly sum- Additions to the Ninth Edition
marizes how each theme was developed in that chapter. Four new or expanded features appear in the Ninth
The four major themes provide excellent topics for essay Edition of Biopsychology.
assignments and exam questions.
NEW! Chapter-Opening Study Objectives Each
chapter begins with a list of study objectives designed to
direct students’ reading and studying.
xvi
NEW! Blog-On! Biopsychology now comes with an Chapter 4: Neural Conduction and Synaptic
accompanying author-run blog and website (www.biopsyc. Transmission
com). The blog contains discussions of exciting new bio-
Simplified explanation of the resting potential
psychological research and theoretical issues not covered
•
Recent discovery that action potentials originate in
in the text. In addition to the blog, the website also con-
•
the axon initial segment
tains a wide variety of links and materials to help students
• Role of glia in synaptic transmission
in their studying.
• Additional coverage of gap junctions
• 22 new citations
NEW! Even More MyPsychLab (www.mypsychlab.
com) MyPsychLab is an online study resource that offers Chapter 5: The Research Methods of
a wealth of animations and practice tests as well as addi- Biopsychology
tional study and research tools. This edition adds even more
Use of PET to determine the distribution in the brain
exciting content to MyPsychLab for students. •
of particular molecules
• Introduction of diffusion tensor imaging with a new
NEW! More Illustrations and Brain Images illustration
Building on Biopsychology’s strong art package, a number
• Simplified coverage of reversible brain lesions
of new illustrations and brain images have been added.
• Updated coverage of gene replacement
These have been carefully selected, designed, and posi-
• 19 new citations
tioned to support interest, clarity, and memorability.
Chapter 6: The Visual System
New Coverage in the Ninth Edition • Simplified coverage of the organization of neurons in
the primary visual cortex
Biopsychology remains one of the most rapidly progress-
• Explanation of the role of context on the responses of
ing scientific fields. Like previous editions, the Ninth visual neurons
Edition of Biopsychology has meticulously incorporated
• Improved discussion of dorsal and ventral streams
recent developments in the field—it contains more than
• Comparison of MT and fusiform face area
750 citations of articles or books that did not appear in
• 24 new citations
the preceding edition. These recent developments have
dictated changes to many parts of the text. The following Chapter 7: Mechanisms of Perception: Hearing,
list presents some of the content changes to this edition, Touch, Smell, Taste, and Attention
organized by chapter.
• Updated coverage of primary auditory cortex
• Statement of the role of cutaneous receptors in par-
Chapter 1: Biopsychology as a Neuroscience
ticular sensations
• 3 new citations • More comparisons of auditory and visual cortex
Chapter 2: Evolution, Genetics, and Experience • Improved coverage of the olfactory system
Chapter 3: Anatomy of the Nervous System Chapter 9: Development of the Nervous System
• Updated coverage of glial function • Updated description of increased cell fate specifica-
• 15 new citations tion and related stem cell terminology
• New description of interaction of glial and neural • Circadian activity cycles of SCN neurons
development • 33 new citations
• Discovery that radial glial cells develop into neurons
• Overall editing designed to shorten and simplify Chapter 15: Drug Addiction and the Brain’s
• 59 new citations Reward Circuits
Chapter 10: Brain Damage and Neuroplasticity • International statistics of drug use
• Increased coverage of the therapeutic effects of THC
• Definition of gliomas • Increased coverage of the effects of MDMA
• Improved coverage of strokes • Updated description of the treatment of heroin
• Updated discussion of genetic factors and neurologi-
addiction
cal disorders • Section on critical thinking about illegal drugs
• Introduction of the term absence seizures
• Major revision of discussion of early theories of
• More concise coverage of the etiology of multiple
addiction
sclerosis • Major update of current approaches to the study of
• Concise summary of genes linked to Alzheimer’s disease
addiction
• Role of microbleeds in Alzheimer’s disease
• 102 new citations
• Importance of early diagnosis in the treatment of
Alzheimer’s disease
Chapter 16: Lateralization, Language, and the
• Improved coverage of MPTP model
Split Brain
• Revised coverage of recovery of function
• Neurotransplantation is revised, updated, and out in • Discussion of the interactions between the hemi-
a historical perspective spheres of split brains
• 95 new citations • Current status of the theory that right hemispheres
are specialized for emotion
Chapter 11: Learning, Memory, and Amnesia • Update of research on neuroanatomical asymmetries
• 45 new citations
Chapter 13: Hormones and Sex
• New section on the modern perspective of sex differ- Chapter 18: Biopsychology of Psychiatric
ences in the brain Disorders
• New section on development of sex differences in hu-
• Introduction of the anticipated DSM-V and the need
man behavior for continual refinement of diagnoses
• Discussion of sex differences in susceptibility to disease
• New section on the current research and treatment of
• Updated coverage of anabolic steroid use
schizophrenia
• New coverage of human sexual arousal and the brain
• Updated coverage of treatments for depression
• 56 new citations
• Updated discussion of the monoamine theory of
depression
Chapter 14: Sleep, Dreaming, and Circadian
• New description of the neuroplasticity theory of
Rhythms
depression
• Updated coverage of the relationship between REM • Discussion of current treatments for anxiety
sleep and dreaming • Discussion of problems with current system of diagnosis
• Introduction to the idea that pharmaceutical compa- • Key Terms appear in boldface, and other important
nies suppress negative findings terms of lesser significance appear in italics.
• 60 new research citations
• Appendixes serve as convenient sources of additional
information for students who want to expand their
Pedagogical Learning Aids knowledge of selected biopsychology topics.
Biopsychology has several features expressly designed to
help students learn and remember the material: Ancillary Materials Available with
• Scan Your Brain study exercises appear within chap- Biopsychology
ters at key transition points, where students can ben-
efit most from pausing to consolidate material before For Instructors
continuing. Pearson Education is pleased to offer the following sup-
plements to qualified adopters.
1. _______________________ lobe beled with a topic and a page reference so that instruc-
2. _______________________ gyrus
3. _______________________
tors can easily select appropriate questions for their tests.
4. _______________________ Textbook authors rarely prepare their own test banks;
5. _______________________ the fact that Pinel insists on preparing the Biopsychology
6. _______________________
7. _______________________ colliculus
test bank attests to its consistency with the text—and his
8. _______________________ body commitment to helping students learn.
9. _______________________
14. _______________________
For Students
MyPsychLab (0205988288) With this exciting new tool,
students are able to self-assess using embedded diagnos-
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• Customizable—MyPsychLab is customizable. in- The customized study plan will focus on the student’s
structors choose what students’ courses looks like. strengths and weaknesses, based on the results of the
Homework, applications, and more can easily be diagnostic testing, and present a list of activities and re-
turned on and off. sources for review and remediation, organized by chap-
• Blackboard Single Sign-on—MyPsychLab can be ter section. Some study resources intended for use with
used by itself or linked to any course management portable electronic devices, such as key terms flashcards
system. Blackboard single sign-on provides deep link- and video clips, are made available exclusively through
ing to all New MyPsychLab resources. MyPsychLab. Students will be able to quickly and eas-
• Pearson eText and Chapter Audio—Like the printed ily analyze their own comprehension level of the course
text, students can highlight relevant passages and add material and study more efficiently, leading to excep-
notes. The Pearson eText can be accessed through tional exam results! An access code is required and can
laptops, iPads, and tablets. Download the free Pearson be purchased at www.pearsonhighered.com or at www
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plans promote better critical thinking skills. The study
plan organizes students’ study needs into sections A Colorful Introduction to the Anatomy of the Human
such as Remembering, Understanding, Applying, and Brain, Second Edition (0205548741) This book, written
Analyzing. by John P. J. Pinel and Maggie Edwards, provides an easy
• MyPsychLab Margin Icons—Margin icons guide and enjoyable means of learning or reviewing the funda-
students from their reading material to relevant vid- mentals of human neuroanatomy through the acclaimed
eos and simulations. directed-coloring method.
Patti Simone, Santa Clara University Dennis Vincenzi, University of Central Florida
Ken Sobel, University of Central Arkansas Ashkat Vyas, Hunter College
David Soderquist, University of North Carolina at Charles Weaver, Baylor University
Greensboro Linda Walsh, University of Northern Iowa
Michael Stoloff, James Madison University David Widman, Juniata College
Stuart Tousman, Rockford College Jon Williams, Kenyon College
Dallas Treit, University of Alberta David Yager, University of Maryland
Margaret Upchurch, Transylvania University H.P. Ziegler, Hunter College
I
n the 1960s, I was, in the parlance of the times, “turned into conventional “textbookese,” and it never let me for-
on” by an undergraduate course in biopsychology. I get that I was writing this book for you.
could not imagine anything more interesting than a Creative thinking is one of the major themes of this
field of science dedicated to studying the relation between edition. Often science and creativity are considered to
psychological processes and the brain. My initial fascina- be opposites, but in my experience many of the major
tion led to a long career as a student, researcher, teacher, advances in biopsychological science have resulted from
and writer of biopsychological science. Biopsychology is creative thinking. These major advances have been made
my attempt to share my fascination with you. by biopsychologists who have recognized that there are
I have tried to make Biopsychology a different kind alternatives to the conventional ways of thinking about
of textbook, a textbook that includes clear, concise, and biopsychological issues that have been engrained in them
well-organized explanations of the key points but is still by their culture and training and who have adopted cre-
interesting to read—a book from which you might sug- ative new approaches. Two things in particular have fas-
gest suitable sections to an interested friend or relative. cinated me about the interplay between creative thinking
To accomplish this goal, I thought about what kind of and biopsychological science: how difficult it is to iden-
textbook I would have liked when I was a student, and I tify and shed conventional approaches even when they
decided to avoid the stern formality and ponderous style clearly haven’t been working, and how often solutions
of conventional textbook writing and to focus on ideas of to long-standing problems become apparent when ap-
relevance to your personal life. proached from a new perspective. The focus of this edi-
I wanted Biopsychology to have a relaxed and personal tion on creative thinking is intended to make the study of
style. In order to accomplish this, I imagined that you biopsychology more interesting for you and to encourage
and I were chatting as I wrote, and that I was telling you become a more creative thinker.
you—usually over a glass of something—about the in- I hope that Biopsychology teaches you much of rele-
teresting things that go on in the field of biopsychology. vance to your personal life and that reading it generates
Imagining these chats kept my writing from drifting back in you the same positive feelings that writing it did in me.
J
ohn Pinel, the author of Biopsychology, obtained Pinel attributes much of his success to his wife,
his Ph.D. from McGill University in Montreal Maggie, who has at various times been a professional
and worked briefly at the Massachusetts Institute artist, designer, and personal trainer. Over the years, they
of Technology before taking a faculty position at the have collaborated on many projects, and the high quality
University of British Columbia in Vancouver, where he of Biopsychology’s illustrations is largely attributable to
is currently Professor Emeritus. Professor Pinel is an her skill and effort.
award-winning teacher and the author of more than 200 Pinel is an enthusiastic West African drummer who
scientific papers. However, he feels that Biopsychology is performs at local clubs, festivals, and drum circles with
his major career-related accomplishment: “It ties together Nigerian drum master Kwasi Iruoje. For relaxation, he
everything I love about my job: students, teaching, writ- loves to cuddle his three cats: Rastaman, Sambala, and
ing, and research.” Squeak.
xxiii